UNIVERSITY  OF  CALIFORNIA   PUBLICATIONS 

IN 

AGRICULTURAL   SCIENCES 

Vol.  3,  No.  8,  pp.  135-242,  plates  13-24  July  12,  1918 


THE  CHEMICAL  COMPOSITION  OF  THE  PLANT 
AS  FURTHER  PROOF  OF  THE  CLOSE  RELA- 
TION BETWEEN  ANTAGONISM  AND 
CELL  PERMEABILITY 

BY 

DEAN  DAVID  WAYNICK 


CONTENTS 

PAGE 

Introduction    , 135 

Object  of  the  investigation 137 

Review  of  previous  investigations  137 

Methods    140 

Experimental  data  144 

External  appearances  of  the  plants 154 

General  review  of  experimental  results 155 

Results  with  salts  of  the  heavy  metals 156 

Possible  effects  of  variations  in  the  concentrations  of  the   solutions  on  the 

plants 160 

Consideration  of  a  possible  Calcium-Magnesium  ratio  160 

Permeability  and  antagonism 162 

Summary    164 


Introduction 

A  solution  of  a  single  salt  at  certain  concentrations  is  toxic  to 
plants  grown  in  it.  The  addition  of  a  second  salt  usually  permits  of 
growth  superior  to  that  in  a  solution  of  a  single  salt  alone  even  though 
the  added  salt  is  toxic  when  used  by  itself.  A  third  salt  added  may 
permit  of  a  still  further  increase  over  the  growth  in  the  two  salt  solu- 
tion. Other  salts  added  will  increase  or  decrease  growth,  depending 
upon  the  salt  used.  Qualitative  relationships  only  have  been  consid- 
ered. When  we  adjust  the  quantitative  relationships  of  the  various 
salts  present,  having  at  the  same  time  due  regard  for  their  qualitative 


136  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

nature,  we  get  as  a  result  a  solution  in  which  the  plant  grows  and 
functions  normally.  Such  a  solution  has  been  termed  by  Loeb,  "phy- 
siologically balanced. ' ' 

It  is  evident  that  if  growth  is  better  in  a  two  salt  solution  the  toxic 
effects  of  the  solution  due  to  a  single  salt  must  be  lessened  by  the 
presence  of  the  second  salt.  We  may  refer  to  either  as  the  second  salt 
since  either  may  be  toxic  alone.  On  the  addition  of  a  third  salt  the 
increase  in  growth  over  that  obtained  in  the  two  salt  solution  points 
to  a  still  further  lessening  of  the  toxic  properties  of  the  various  salts 
present  taken  singly.  This  action  of  one  or  more  salts  in  limiting  or 
preventing  entirely  the  toxic  effects  of  one  or  more  other  salts,  is 
termed  antagonism.  Sea  water  may  be  taken  as  an  example  of  a 
physiologically  balanced  solution  or  a  solution  in  which  the  mutual 
antagonism  between  the  constituents  of  the  solution  is  such  as  to 
allow  of  normal  growth  of  numerous  organisms. 

The  fact  of  the  existence  of  antagonism  has  been  proven  by  a 
number  of  investigators  working  in  plant  and  animal  physiology,  but 
the  mechanism  of  antagonistic  action  is  by  no  means  clear.  Since 
salts  are  very  largely  ionized  in  the  nutrient  solutions  usually  em- 
ployed, it  is  probable  that  antagonism  has  to  do  with  ions.  Further, 
antagonism  will  probably  take  place  between  the  ions  present  in,  or 
between,  the  ionic  constituents  of  the  solution,  and  the  living  mem- 
branes in  contact  with  the  solution.  Loeb1  first  advanced  the  theory 
that  one  ion  may  prevent  the  entrance  of  another  ion  into  living  cells 
and  that  in  this  property  lies  the  reason  for  antagonistic  action.  On 
the  basis  of  this  hypothesis,  penetration  precedes  the  manifestations  of 
toxic  effects  and  where  penetration  does  not  occur,  due  to  antagonistic 
action,  there  are  no  toxic  effects  evident.  Used  in  this  way,  the  term 
penetration  means  simply  the  entrance  of  ions  in  greater  number  than 
would  normally  occur  were  the  plant  cells  in  their  natural  environ- 
ment. Experimental  evidence  as  to  the  correctness  of  this  hypothesis 
has  been  furnished  by  Loeb2  in  a  very  interesting  series  of  experi- 
ments. Ost^rhout3  has  applied  the  electrical  conductivity  method 
to  the  measurement  of  the  penetration  of  ions  into  plant  tissue, 
while  recently  Brooks  has  confirmed  Osterh out's  results  (1)  by  deter- 
mining: the  diffusion  of  ions  through  tissue,4  (2)  by  exosmosis,5  and 
(3)  by  the  change  in  the  curvature  of  tissue.0 


i  Amor.  Jour.  Physiol.,  vol.  6  (1902),  p.  411. 

-Science,  n.s.,    vol.  36,  no.  932,  p.   637. 

a  Ibid.,  vol.  35,  no.  890,  p.  112. 

'  Proc.    Nat.   Acad.,  Sci.,  vol.  2   (1916),  p.  569. 

■Anier.  .lour.    Hot.,   vol.  3    (1916),  p.  483. 


1918]  Waynick:    Antagonism  and  Cell  Permeability  137 

It  is  evident  that  these  methods  are  limited  in  their  application 
and  give  no  idea  of  the  quantitative  relationships  existing  between 
the  ions  actually  entering  the  cells.  They  do  show,  however,  that  the 
permeability  of  the  plant  tissue  may  be  greatly  altered  by  salt  action 
and  that  solutions  which  permit  of  normal  growth,  also  preserve  normal 
permeability  as  regards  the  ions  present  in  the  solution. 

Object  of  the  Investigation 

In  a  preliminary  paper7  the  results  obtained  from  chemical  analy- 
ses of  plants  grown  in  toxic  and  antagonistic  solutions  have  been 
reported.  These  results  were  of  interest  and  the  general  method  em- 
ployed seemed  to  be  worthy  of  a  more  extended  application  in  the 
determination  of  ions  absorbed  by  plants  from  solutions,  of  known 
composition  and  concentration.  From  a  consideration  of  the  data 
in  the  paper  referred  to  above,  it  was  felt  that  the  results  obtained  in 
a  more  extensive  investigation  would  be  of  importance:  (1)  from 
the  standpoint  of  the  effect  of  various  salts  upon  the  permeability  of 
the  cell  tissue  of  growing  plants;  (2)  from  that  of  the  effects  of  vari- 
ous salts  upon  the  nutrition  of  plants  as  evidenced  by  growth;  (3) 
from  that  of  a  possible  correlation  of  growth  with  the  absorption  of 
ions;  and  (4)  from  the  standpoint  of  the  quantitative  relationships 
existing  between  certain  ions  of  the  solution  and  the  same  ionic  rela- 
tionships in  the  plant. 

The  various  phases  of  the  problem  as  outlined  above  will  be  con- 
sidered in  the  discussion  of  the  experimental  results  following. 

Review  of  Previous  Investigations 

It  is  not  intended  that  the  following  review  of  the  previous  work 
done  in  this  field  of  plant  physiology  be  exhaustive.  Robertson8  has 
reviewed  the  literature  dealing  with  antagonistic  salt  action  very 
completely  up  to  a  recent  date.  Brenchley9  and  Lipman  and  Gericke10 
have  referred  to  all  the  important  work  done  with  regard  to  the 
effects  of  the  salts  of  the  heavy  metals  upon  plants.  The  present 
review  therefore  touches  only  the   work  bearing   directly  •  upon  the 


a  Ibid.,  p.  562. 

'  Contribution  to  the  causes  of  antagonism  between  ions.  (Univ.  Calif., 
Master's  thesis,  1915.) 

sErgeb.  Physiol.  Jahrb.,  vol.  10   (1910),  p.  216. 

9  Inorganic  plant  poisons  and  stimulants.  New  York,  Putnam,  1915  (Cam- 
bridge agricultural  monographs). 

i"Univ.  Calif.  Publ.  Agr.  Sci.,  vol.  1  (1917),  p.  495. 


138  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

present  problem  or  work  so  recent  as  not  to  be  included  in  the  papers 
cited  above. 

A  large  share  of  the  contribution  to  the  experimental  evidence  in 
regard  to  antagonism  between  salts  as  regards  plants  we  owe  to  Oster- 
hout.  In  a  series  of  papers  he  has  shown  that  any  salt  may  be  toxic 
to  plants  when  used  alone  in  solution  at  certain  concentrations  and 
further  that  the  addition  of  a  second  salt  may,  in  proper  concentra- 
tion, modify  or  eliminate  entirely  the  toxic  effect  of  the  first  salt.  He 
has  shown  further  that  acids,  alkalies,  and  various  organic  compounds 
may  likewise  be  toxic  to  plants  and  that  their  toxic  effects  may  be 
modified  by  the  presence  of  a  variety  of  compounds,  depending  upon 
the  toxic  substance  employed.  By  measuring  the  resistance  of  cylin- 
ders of  Laminaria  in  solutions  of  one  salt  and  in  solutions  containing 
two  or  more  salts,  he  has  brought  forward  much  evidence  as  to  the 
penetration  of  ions  into  plant  cells.  While  this  method  has  yielded 
very  valuable  results  both  as  to  the  rate  of  entrance  of  ions  and  also 
the  total  number  of  ions  penetrating,  it  does  not  yield  results  which 
give  us  a  knowledge  of  the  relative  amounts  of  the  various  ions  which 
penetrate  the  tissue  when  the  qualitative  as  well  as  the  quantitative 
relationships  of  the  nutrient  solution  are  varied.  Osterhout  has 
shown,  however,  that  penetration  is  more  rapid,  and  the  degree  of 
permeability  is  greatly  increased,  in  unbalanced  solutions  and  further 
that  as  the  permeability  of  the  plant  tissue  more  nearly  approaches 
normal  the  growth  of  the  plant  is  also  more  nearly  normal. 

Szucs11  has  used  Cucurbit  a  pepo  as  an  indicator  by  immersing  the 
young  seedlings  in  various  solutions  for  varying  periods  of  time  and 
counting  those  still  able  to  show  geotropic  movement  when  placed  in  a 
horizontal  position  in  a  moist  chamber.  He  found  a  marked  antagon- 
ism between  copper  sulphate  and  aluminum  chloride  and  concludes 
from  his  experiments  that  antagonism  consists  in  the  mutual  hin- 
drance of  similarly  charged  ions  in  entering  the  cell.  He  states 
further  that  the  rate  of  absorption  of  equally  charged  ions  is  of  great 
importance.  His  chemical  methods  are  open  to  question,  for  in  the 
experiments  reported  the  test  for  copper  used  was  that  of  boiling  the 
roots  and  testing  the  resulting  solution  for  copper  with  hydrogen 
sulphide. 

By  analyzing  the  solution  in  which  pea  seedlings  had  grown,  Pan- 
ic! li'-  has  determined  ion  absorption.     The  growing  period  was  short. 


ujahrb.  Wies.  Bot.  (Pringheim) ,  vol.  52,  no.  1  (1912),  p.  85. 
>-  Ibid.,  p.  211. 


1918]  Waynick:    Antagonism  and  Cell  Permeability  139 

He  found  a  rapid  absorption  of  zinc,  manganese,  iron,  and  aluminum, 
but  the  total  amounts  taken  up  were  small.  He  gives  other  evidence 
of  the  selective  absorption  of  various  other  ions  from  solutions,  but 
these  results  are  of  not  direct  application  here.  It  is  of  interest  to 
note,  however,  that  he  found  a  direct  relation  between  time  and  ion 
absorption.  His  most  important  conclusion,  which  bears  directly  upon 
the  problem  in  hand,  is  that  strong  narcosis  was  associated  with  the 
penetration  of  ions  in  large  numbers. 

Schreiner  and  Skinner,13  using  a  similar  method,  have  determined 
the  amounts  of  phosphoric  acid,  nitrates,  and  potassium  remaining  in 
a  solution  in  which  plants  had  been  grown.  Various  ratios  of  these 
three  ions  were  employed,  the  total  concentration  being  80  parts  per 
million.  They  found  widely  varying  amounts  of  these  three  ions 
removed  from  the  solution,  and  further  there  seemed  to  be  a  possible 
difference  of  20  to  30  per  cent  in  the  removal  of  any  one  without 
an  apparent  effect  upon  the  growth  of  the  plants.  Under  the  condi- 
tions reported  by  them  increased  growth  was  correlated  with  increased 
absorption. 

By  means  of  conductivity  measurements  of  solutions  in  which  pea 
seedlings  were  growing,  True  and  Bartlett14,  15'  16  have  determined 
the  rate  of  absorption  and  of  excretion  of  electrolytes.  Their  work 
was  done  with  one,  two  and  three  salt  solutions.  In  general  they 
found  a  greater  absorption  when  a  mixture  of  salts  was  present  than 
when  single  salts  were  used.  Further,  the  absorption  relationships 
of  salts  with  a  common  kation  seem  to  be  similar.  For  example,  from 
solutions  of  low  concentrations,  potassium  chloride,  potassium  sul- 
phate, and  potassium  nitrate  are  not  removed,  but  on  the  other  hand 
there  is  an  excretion  of  electrolytes  by  the  plant.  In  direct  contrast, 
calcium  nitrate  and  calcium  sulphate  are  removed  from  their  solu- 
tions in  every  concentration  employed  and  no  excretion  of  electro- 
lytes from  the  plants  could  be  detected.  It  seems  probable  that  the 
low  concentration  employed  by  them  acted  as  a  limiting  factor  in 
some  cases. 

In  a  recent  paper  Breazeale17  has  shown  that  the  presence  of 
sodium  carbonate,  and  sodium  sulphate,  when  used  in  concentrations 
of  1000  parts  per  million  in  nutrient  solutions,  decreased  the  absorp- 


isBot.  Gaz.,  vol.  50  (1910),  p.  1. 

HAmer.  Jour.  Bot,  vol.  2   (1915),  p.  255. 

is  Ibid.,  p.  311. 

^  Ibid.,  vol.  3  (1915),  p.  47. 

17  Jour.  Agr.  Kesearch,  vol.  7  (1916),  p.  407. 


140  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

tion  of  potassium  and  phosphoric  acid  as  much  as  70  per  cent  below 
that  of  the  control  cultures. 

The  work  of  Gile18  is  of  interest  in  this  connection.  From  ash 
analyses  obtained  in  investigating  the  cause  of  chlorosis  in  pineapples, 
he  found  a  direct  relationship  between  the  absorption  of  lime  and 
that  of  iron;  that  is,  when  the  absorption  of  lime  was  high  but  little 
iron  was  taken  up.  In  soil  studies  Gile  and  Ageton19  found  no  direct 
relation  between  the  lime  content  of  plants  and  varying  amounts  of 
lime  and  magnesia  in  the  soil. 

A  few  investigations  have  been  made  on  the  absorption  of  specific 
elements  from  solution,  but  these  need  only  be  mentioned  in  the 
present  connection.  Maquenne20  found  that  mercuric  chloride  causes 
marked  increase  in  permeability  of  the  protoplasm,  although  it  is  not 
necessarily  absorbed  itself  in  any  considerable  quantities.  Marsh21 
correlates  the  amount  of  barium  chloride  present  in  the  soil  with 
that  found  in  the  plant.  Colin  and  De  Rufz22  always  found  absorbed 
barium  localized  in  the  roots. 

A  large  number  of  analyses  of  plants  grown  under  various  condi- 
tions have  been  reported,  but  the  environmental  factors  have  varied 
so  greatly  as  to  render  the  results  obtained  of  little  value  in  the 
present  study. 

From  this  review  it  is  evident  that  no  quantitative  study  of  the 
elements  actually  absorbed 'from  the  nutrient  solutions,  balanced  and 
unbalanced,  has  been  made  with  the  idea  in  mind  of  a  correlation 
between  the  absorption  of  the  various  ions  with  their  antagonistic 
or  toxic  effects  in  solution  cultures. 

Methods 

Barley  was  used  as  the  plant  indicator.  The  seeds  were  obtained 
from  the  University  Farm  at  Davis  and  were  of  a  pure  strain  of  the 
Beldi  variety.  The  method  of  sprouting  the  seeds,  while  simple,  has 
not  been  noted  elsewhere  and  has  given  such  excellent  results,  both  to 
the  writer  and  to  others,  that  it  seems  worthy  of  mention  here  in 
detail.  A  piece  of  oilcloth  about  12x18  inches  was  covered  with  sev- 
eral thicknesses  of  paper  toweling  and  the  whole  thoroughly  wetted. 


is  I'orto  Rico  Exp.  Sta.  Bull.,  11    (1911). 

ifl  Ibid.,   Bull.  10  (11)14). 

20C.-R.  Acad.  Sci.  (Paris),  vol.  123  (1896),  p.  898. 

2i  Bot.  Gaz.,  vol.  54   (1912),  p.  2~>0. 

22C.  B.  Acad.  Sci.   (Paris),  vol.  150  (1910),  p.  1074. 


1918]  Waynick:    Antagonism  and  Cell  Permeability  141 

Selected  seeds  were  distributed  over  the  toweling  so  that  about  two 
hundred  were  placed  on  an  area  of  the  size  indicated  above.  Another 
layer,  made  up  of  several  sheets  of  toweling-,  was  then  laid  on  the 
seeds  and  the  whole  thoroughly  soaked  with  water.  The  water  was 
allowed  to  evaporate  gradually  until  the  paper  was  but  slightly  moist 
to  the  touch  and  the  water  relation  then  maintained  constant  until 
the  seedlings  were  transferred  to  the  solutions.  If  the  paper  is  kept 
too  moist  the  growth  of  molds  is  often  very  abundant,  but  with  a  low 
moisture  content  no  trouble  was  experienced  from  this  source.  By  the 
time  the  roots  were  a  quarter  of  an  inch  long,  the  upper  layer  of 
paper  was  supported  two  or  three  inches  above  the  seedlings.  This 
procedure  permits  of  a  straight  growth  of  the  shoots,  which  is  of  con- 
siderable importance  in  placing  the  seedlings  in  the  corks.  The  seed- 
lings were  transferred  when  the  shoots  were  about  an  inch  and  a  half 
in  length.  The  paper  in  which  the  roots  are  grown,  tears  apart  readily 
Avithout  injuring  them  in  any  way,  the  oilcloth  not  permitting  their 
downward  penetration.  There  is  no  contact  with  metal  containers 
at  any  time,  the  apparatus  required  is  practically  nothing,  the  time 
period  is  short — about  six  days  under  greenhouse  conditions — and 
strong  seedlings  are  obtained  which  can  be  transferred  to  any  contain- 
ers without  injury. 

The  containers  used  were  quart  jars  of  the  Mason  type,  each 
holding  approximately  950  c.c.  of  solution.  The  inside  of  each  jar, 
as  well  as  that  of  the  bottles  for  the  stock  solutions,  was  coated  with 
a  layer  of  paraffin  so  that  the  solutions  were  never  in  contact  with  the 
glass.  The  outside  of  the  jar  was  covered  with  black  paper  to  exclude 
light,  the  black  surface  facing  the  glass.  Flat  corks,  having  a  diam- 
eter of  three  and  a  half  inches,  were  used  to  support  the  seedlings. 
Each  cork  had  seven  holes,  one  in  the  center  through  which  distilled 
water  was  added  to  maintain  the  volume  of  the  solution  as  nearly 
constant  as  possible,  and  six  equally  spaced,  one  and  a  quarter 
inches  from  the  center,  for  holding  the  seedlings.  After  the  holes 
were  made  the  corks  were  soaked  in  boiling  paraffin. 

To  introduce  the  seedlings  the  corks  were  turned  upside  down, 
supported  by  the  rim  of  the  jar,  and  the  shoots  stuck  through  the 
holes  prepared  for  them  and  held  in  place  by  a  small  piece  of  cotton. 
On  turning  the  corks  over  the  seedlings  were  in  their  proper  position 
without  being  in  the  least  injured,  for  there  was  no  necessity  for 
touching  the  roots  at  any  stage  since  the  plant  was  always  picked  up 
by  the  seed  coat.     The  method  suggested  by  Tottingham23  was  tried, 


142  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

but    the    one    outlined    above    proved    very    satisfactory    and    much 
simpler. 

The  basic  nutrient  solution  used  throughout  was  Shive's  three 
salt  nutrient24  containing  the  following  salts  in  the  given  partial 
molecular  concentrations : 

K  H2  P04  0180  M. 

Ca   (N03)2 0052  M. 

MgS04  0150  M. 

The  stock  solution  was  made  up  to  twice  the  strength  indicated 
above  and  diluted  as  necessary  by  the  addition  of  added  salt  solution, 
or  distilled  water,  or  both. 

In  the  case  of  the  chlorides  used,  viz.,  calcium,  magnesium  and 
potassium,  normal  or  twice  normal  solutions  were  prepared  and 
standardized  by  titrating  against  a  standard  silver  nitrate  solution. 
Normal  solutions  of  magnesium  and  potassium  sulphate  were  stand- 
ardized by  weighing  the  barium  sulphate  precipitate.  Solutions  of 
copper,  zinc,  iron,  and  mercury  salts  were  prepared  in  concentrations 
of  1000  parts  per  million  by  weighing  out  the  carefully  dried  salts. 

The  final  volume  of  solution  required  for  the  duplicate  jars  was 
approximately  two  thousand  cubic  centimenters.  Starting  with  a 
thousand  of  the  nutrient  solution,  various  volumes  of  the  standard 
solutions  were  added  so  that  when  the  total  volume  was  made  up  to 
two  liters  with  distilled  water,  the  concentrations  of  the  various  salts 
would  be  those  reported  in  the  accompanying  tables. 

The  growing  period  was  six  weeks.  The  duplicate  cultures  were 
grown  in  specially  constructed  mouse-proof  cages  each  holding  ninety 
jars.  The  tops  of  the  cages  were  open  and  the  sides  made  of  coarse 
wire  screening.  The  different  parts  of  the  cages  were  equally  well 
lighted,  as  shown  by  the  nearly  equal  growth  of  the  controls  in  dif- 
ferent parts  of  the  cages.  When  necessary  the  plants  were  supported 
by  cords  strung  across  from  side  to  side  of  the  cages. 

The  solutions  were  not  changed  during  the  growing  period,  but 
the  volumes  were  kept  as  nearly  constant  as  possible  by  adding  dis- 
tilled water.  There  are  objections  to  this  method,  as  there  are  objec- 
tions to  the  method  of  using  water  cultures  at  all.  The  growth  was 
found  to  be  very  satisfactory  and  compares  favorably  with  the  growth 


28  Physiol.  Researches,  no.  4  (1915),  p.  174. 
24  Arner.  Jour.  Bot.,  vol.  2   (1915),  p.  157. 


1918]  Waynick:   Antagonism  and  Cell  Permeability  143 

obtained  by  other  investigators  in  comparable  periods  of  time.  A 
further  discussion  of  this  point  will  be  taken  np  below. 

At  the  expiration  of  the  six  weeks  growing  period  the  plants 
were  removed  from  the  corks,  the  roots  rinsed  thoroughly  with  dis- 
tilled water,  placed  between  layers  of  paper  toweling,  dried  in  the 
oven  at  100°-105?C,  roots  and  tops  separated,  weighed,  and  placed 
in  envelopes  ready  for  analysis.  For  analysis  the  roots  from  dupli- 
cate cultures  were  combined  unless  the  dry  weight  was  sufficient  to 
allow  of  separate  analysis. 

Total  ash  was  determined  after  direct  ignition  of  the  dry  material 
in  a  muffle  at  a  low  red  heat  until  no  trace  of  carbon  remained.  The 
ash  was  then  taken  up  in  dilute  hydrochloric  acid  and  evaporated 

to  dryness  to  remove  possible  contamination  with  silica.      Iron  was 

N 
precipitated  as  the  hydroxide  with  ammonia  and  titrated  with  — — 

potassium  permanganate  after  reduction  with  zinc  and  sulphuric  acid. 
This  determination  was  made  because  of  the  relation  Gile  has  shown 

to  exist  between  calcium  and  iron  absorption  by  plants.     Calcium 

N 
was  precipitated  as  oxalate  and  titrated  with  — —  potassium  perman- 
ganate. The  double  precipitation  of  the  oxalate  assured  freedom 
from  magnesium  contamination.  Magnesium  was  precipitated  by 
ammonium  phosphate  and  weighed  as  the  pyrophosphate.  Potassium, 
where  determined,  was  precipitated  and  weighed  as  the  chloroplati- 
nate.  Copper  was  determined  colorometrically  by  using  the  ferro- 
cyanide  method.  The  amount  of  material  available  precluded  the 
possibility  of  a  more  complete  analysis  than  was  made  if  any  degree 
of  accuracy  was  desired.  For  example,  in  Series  vn,  the  weight  of 
the  ash  varied  from  12  to  233  milligrams  in  the  case  of  the  roots  and 
from  32  to  183  milligrams  in  the  case  of  the  tops.  While  these  varia- 
tions are  not  extreme,  they  are  fairly  representative.  The  values  of 
these  elements  actually  determined  cannot  be  taken  as  absolute  in 
every  case  because  of  the  limited  amounts  of  material  available,  but 
the  significant  differences  are  so  great  as  to  make  a  small  variation 
in  this  regard  of  minor  importance. 

The  strength  of  all  solutions  is  uniformly  expressed  in  terms  of 
molecular  concentrations  since  this  mode  of  expression  has  been 
quite  generally  used  in  experimental  work  reported  by  different 
investigators. 

Under  experimental  results  twenty-six  series  are  reported.  A 
series,  as  used  in  the  present  work,  may  be  defined  as  a  number  of 


144  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

duplicate  cultures  containing  one  salt  in  varying  concentrations  in 
each,  or  one  salt  constant  and  varying  concentrations  of  a  second  salt. 
In  some  instances  both  salts  varied  but  only  in  concentration,  the 
same  ratios  being  maintained.  These  are  few.  The  number  of  con- 
centrations reported  vary  from  three  to  fourteen  in  a  series,  depend- 
ing upon  the  salt  used.  Before  two  salts  were  taken  together,  the 
effects  of  each  separately  upon  the  plants  were  determined.  Usually 
this  meant  only  the  establishment  of  the  toxic  limits  of  the  salts  em- 
ployed when  used  in  the  nutrient  solution.  Several  series  of  this  kind 
are  not  reported  here,  as  no  analytical  work  was  done  upon  them. 

Calcium  and  magnesium  salts  were  used  to  a  large  extent  because 
of  the  fact  that  their  kations  can  be  determined  with  less  experi- 
mental error  than  most  other  nutrient  salts  where  the  small  amounts 
of  material  dealt  with  here  are  considered;  also  it  was  of  interest 
to  determine  whether  or  not  there  is  a  lime-magnesia  ratio  for  plants 
grown  under  carefully  controlled  conditions.  Copper,  zinc,  iron,  and 
mercury  salts  were  used  because  of  the  fact  that  their  toxic  and  antag- 
onistic effects  have  not  been  previously  determined  as  regards  absorp- 
tion.   Potassium  chloride  was  the  only  monovalent  salt  used. 

A  longer  growing  period  than  has  usually  been  employed  was  con- 
sidered important.  McGowan,25  in  conducting  experiments  in  pure 
solutions  of  sodium,  potassium  and  calcium  chlorides,  found  growth 
better  in  the  first  two  at  the  end  of  six  days,  but  far  superior  in  a 
solution  of  calcium  chloride  in  twenty-five  days.  In  a  qualitative  way 
the  same  relationships  were  observed  in  the  present  investigation.  It 
seems  reasonable  to  assume  that  the  results  obtained  in  six  weeks  with 
plants  are  more  nearly  representative  of  the  true  effect  of  various 
solutions  than  those  obtained  in  two  or  three  day  periods  or  even  in 
three  week  periods.  But  it  is  not  assumed  that  the  results  herein 
reported  are  the  same  as  those  which  might  be  obtained  were  the 
plants  grown  to  maturity.  It  is  hoped  that  more  data  may  be  pre- 
sented shortly  on  this  point. 

In  the  following  section,  in  which  the  experimental  results  are 
given,  the  time  factor  and  the  basic  nutrient  solutions  are  constants. 

Experimental  Data 
All  analyses  are  reported  as  percentages  of  the  dry  weights  of  the 
plants.     To  make  the  results  obtained  as  clear  as  possible,  graphs  and 
photographs  have  been  used  throughout  as  well  as  the  tables  giving 
the  actual   percentage  composition  of  the  plants. 


25  Hot.   Gaz.,  vol.  45   (1908),  p.  45. 


1918]  Wayhick:    Antagonism  and  Cell  Permeability  145 

The  relationships  of  calcium  to  magnesium  salts  are  reported  in 
the  first  seven  tables.  For  a  review  of  the  more  important  literature 
bearing  directly  upon  the  relationships  of  the  salts  to  these  two  ele- 
ments reference  is  made  to  McCool,26  who  has  considered  these  in 
some  detail,  and  to  a  recent  critical  survey  of  the  lime-magnesia  ratio 
hypothesis  b}^  Lipman.27 

As  is  evident  from  table  1,  calcium  chloride  does  not  become  toxic 
until  present  in  concentration  of  over  .24  M.  Up  to  and  including 
this  concentration  the  growth  seems  to  be  but  little  affected  by  the 
increasing  concentrations  of  the  salt  added.  The  percentage  of  cal- 
cium in  the  plants  shows  no  direct  increase  with  increasing  concen- 
tration of  calcium  chloride  in  the  solution.  The  lowest  percentage  of 
calcium  given  occurs  in  a  concentration  of  .20  M.  calcium  chloride. 

In  table  2  there  is  a  close  parallelism  between  the  growth  of  roots 
and  tops.  Two  low  points  on  the  dry  weight  graph  are  evident,  the 
first  occurring  at  cultures  4  and  5  and  the  second  from  7  to  11.  At 
these  low  points  we  have  a  high  percentage  of  magnesium  in  both 
roots  and  tops,  but  of  calcium  only  in  the  second  low  point.  Calcium 
is  low  where  growth  is  good  in  cultures  2  and  3.  But  the  most  inter- 
esting feature  is  the  decreased  absorption  of  both  elements  at  cul- 
ture 6,  where  there  is  a  distinct  increase  in  dry  weight.  Iron  was 
not  present  in  sufficient  concentration  to  allow  of  titration  until  cul- 
ture 11  is  reached.  It  may  be  stated  here  that  the  iron  determined  is 
limited  to  that  in  the  seed  as  a  maximum,  for  it  was  purposely  ex- 
cluded from  the  solutions  except  where  its  toxic  or  antagonistic  action 
was  under  observation.  In  many  instances  the  titration  of  this  residual 
iron  is  of  interest. 

Table  3  is  a  record  of  one  of  the  most  interesting  and  significant 
series  reported.  The  root  growth  was  so  limited  in  nearly  every  cul- 
ture that  no  attempt  was  made  to  segregate  roots  from  tops  for  sep- 
arate determinations  except  where  the  total  dry  weight  was  so  greatly 
increased  as  in  cultures  6  and  11.  In  the  first  place  we  have  double 
maxima  of  growth,  the  first  in  culture  6  and  the  second  in  11.  The 
total  dry  weight  at  culture  11  is  twice  that  at  6,  but  the  dry  weight 
in  culture  6  amounts  to  a  35  per  cent  increase  over  that  in  culture  7. 
A  direct  inverse  relationship  is  shown  between  total  growth  and  ab- 
sorption at  these  two  high  points ;  the  maximum  growth  in  culture  11  is 
accompanied  by  the  lowest  absorption  of  calcium  and  magnesium. 
The  percentage  of  magnesium  is  low  in  culture  6,  but  that  of  calcium 


26  Cornell  Univ.  Agr.  Exp.  Sta.  Mem.  2  (1913),  p.  127, 

27  Plant  world,  vol.  19  (1916),  p.  83. 


146  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

is  higher  than  in  the  cultures  of  slightly  higher  or  lower  concentra- 
tions. No  explanation  of  the  narrow  ratio  between  these  two  ele- 
ments at  this  point  can  be  offered.  It  is  of  interest  to  note  the  very 
great  increase  in  the  amounts  of  calcium  and  magnesium  found  in 
the  plants  grown  in  concentrations  of  .20  M.  calcium  chloride  alone. 
While  magnesium  chloride  is  constant  throughout  the  series,  the 
amount  of  magnesium  does  not  increase  proportionately  to  that  of 
calcium. 

A  still  higher  concentration  of  magnesium  chloride  was  used  in 
the  series  reported  in  table  4.  The  percentage  of  magnesium  found 
in  the  roots  is  very  high  and  would  indicate  that  it  was  not  entirely 
removed  from  the  roots  by  washing.  In  general  the  percentages  of 
calcium  and  magnesium  found  are  high,  the  calcium  content  increas- 
ing as  the  concentration  of  calcium  chloride  present  in  the  culture, 
but  not  proportionately.  Magnesium  is  lower  at  the  greater  dry 
weights  for  the  tops,  the  decrease  amounting  to  50  per  cent  in  the  case 
of  culture  6. 

Magnesium  sulphate  was  used  alone  in  the  series  reported  in 
table  5.  The  decrease  in  growth  is  nearly  proportional  to  the  increase 
in  concentration  of  the  added  salt.  In  this  series  wTe  have  a  very 
marked  decrease  in  the  percentages  of  calcium  and  magnesium  present 
in  the  roots  without  any  evident  effect  upon  the  growth  of  the  plants, 
especially  that  of  the  tops.  Here  again,  however,  we  have  increased 
absorption  of  calcium  as  the  percentage  of  magnesium  increases, 
even  though  the  concentration  of  the  former  in  the  nutrient  solution 
is  constant.  It  is  of  interest  to  note  that  the  percentages  of  both  ele- 
ments in  the  tops  throughout  this  series  are  low  and  vary  but  little, 
regardless  of  the  increasing  concentration  of  the  nutrient  solution. 

Very  marked  antagonism  between  calcium  chloride  and  magnesium 
sulphate  is  shown  in  table  6.  The  dry  weight  of  the  plants  grown  in  a 
solution  of  magnesium  sulphate  .18  M.  concentration  was  .29  gram, 
but  when  .04  M.  concentration  of  calcium  chloride  was  added  the  aver- 
age dry  weight  was  1.20  grams  and  in  a  concentration  of  .18  M. 
magnesium  sulphate  and  .24  M.  calcium  chloride  the  average  dry 
weight  was  .98  gram.  Between  these  two  concentrations  of  calcium 
chloride  the  dry  weights  recorded  are  uniformly  high.  Correlated 
with  the  rapid  decrease  in  growth,  in  concentrations  of  .24  M.  of  cal- 
cium chloride,  is  the  marked  increase  in  the  percentage  of  both  calcium 
and  magnesium  found  in  the  plants.  The  graphs  representing  the 
amounts  of  these  elements  found  crosses  the  growth  graph  coincident 


191S]  Waynick:   Antagonism  and  Cell  Permeability  147 

with  its  sharp  decline.  The  low  percentage  of  magnesium  is  of  interest 
since  the  concentration  of  the  culture  solution  was  uniformly  high 
with  respect  to  this  ion. 

It  is  striking  that  there  is  a  marked  decrease  in  the  growth  of  roots 
at  the  concentration  which  gave  the  best  growth  of  tops,  and  further 
that  the  percentage  of  calcium  in  the  tops  and  magnesium  in  the 
roots  parallel  this  decrease  in  the  growth  of  the  roots.  A  comparison 
of  the  results  obtained  with  magnesium  sulphate  as  against  those  with 
magnesium  chloride  is  reserved  for  later  discussion. 

In  table  7  we  have  an  opportunity  to  compare  indirectly  anion 
effects,  or  possibly  the  effects  of  combinations  of  the  same  kation  with 
different  anions.  From  preliminary  results  it  seemed  advisable  to  use 
.15  M.  magnesium  sulphate  in  this  series  instead  of  .18  M.  as  used  in 
the  preceding  series,  so  that  the  concentration  of  magnesium  ion  is 
not  equivalent  in  the  two  series.  A  solution  containing  magnesium 
sulphate  .15  M.  plus  calcium  nitrate  .08  M.  proved  highly  toxic,  while 
a  solution  containing  calcium  chloride  of  the  same  concentration  as 
the  nitrate  in  the  above  solution  supported  normal  growth.  It  is 
possible  that  the  difference  is  due  to  the  toxic  action  of  the  nitrate  ion 
on  the  plant  directly.  Tottingham  has  shown  that  the  total  ionization 
of  a  nutrient  solution  was  decreased  10  per  cent  below  the  theoretical 
by  the  addition  of  calcium  nitrate  in  low  concentrations.  It  is  pos- 
sible that  the  ionization  of  some  other  salt  is  repressed  so  that  there 
is  an  actual  lack  of  some  ion  necessary  for  growth.  The  percentage 
of  calcium  found  was  not  high  enough  in  any  case  to  account  for  the 
toxic  effects  shown.  Magnesium  Avas  found  in  extremely  large 
amounts,  9.20  per  cent  in  the  case  of  culture  6,  the  largest  percentage 
recorded  in  any  culture  studied.  Unfortunately  the  series  in  which 
the  toxic  effects  of  calcium  nitrate  alone  were  studied  was  lost,  so  it 
cannot  be  reported  here. 

Potassium  chloride  was  the  only  monovalent  salt  studied,  and  the 
results  are  given  in  tables  8  and  9.  The  growth  shown  in  the  various 
concentrations  of  potassium  chloride  used  was  approximately  the 
same  as  that  found  when  magnesium  sulphate  was  used  alone.  The 
increase  in  the  percentage  of  ash,  as  far  as  the  tops  are  concerned  in 
table  8  is  very  striking.  The  percentage  of  calcium  found  in  the  tops 
and  of  magnesium  found  in  the  roots  remains  practically  constant 
throughout.  The  amount  of  potassium  absorbed  increases  as  the  con- 
centration of  potassium  chloride  in  the  solution  increases  and  in- 
versely as  the  growth  of  the  plants.     The  toxic  effects  due  to  the 


148  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

addition  of  potassium  chloride  to  the  solution  are  much  more  evident 
in  the  tops  than  in  the  roots  with  respect  to  the  increasing  concen- 
trations of  potassium  chloride. 

Using  a  constant  concentration  of  potassium  chloride  of  .18  M., 
which  is  an  increase  of  .02  M.  over  the  highest  concentration  of  that 
salt  reported  in  table  8,  against  varying  concentrations  of  magnesium 
sulphate,  the  results  reported  in  table  9  were  obtained.  There  is  a 
marked  increase  in  total  ash  as  the  concentration  of  the  nutrient  solu- 
tion with  respect  to  magnesium  sulphate  increases.  Parallel  with  this 
increase  is  the  higher  percentage  of  potassium.  The  growth  decreases 
inversely.  Antagonism  between  the  two  salts  is  evident  where  the 
lower  concentrations  of  magnesium  sulphate  were  used.  In  cultures  2 
and  4  of  this  series,  we  have  a  marked  increase  in  growth  over  that  of 
culture  3.  Absorption  is  markedly  lower  at  the  two  high  points  than 
at  the  intermediate  concentration,  where  the  solution  is  evidently 
more  toxic.  The  least  growth  obtained  in  the  series  was  recorded  in 
culture  7,  which  shows  the  highest  absorption  of  all  the  elements 
determined.  In  the  two  higher  concentrations  of  magnesium  sulphate 
used  the  growth  was  increased  somewhat  while  the  percentage  of  cal- 
cium, magnesium,  and  potassium  in  the  plants  decreased  markedly. 
It  seems  worthy  of  note  that  the  amount  of  iron  in  the  ash  was  not 
sufficient  to  allow  of  titration  at  any  concentration  employed  in  the 
series.  This  series  very  well  illustrates  the  point  which  has  been 
brought  out  a  number  of  times  before  of  the  relationship  between 
absorption  and  growth.  Here  we  have  five  cultures  in  the  one  series 
of  which  this  relationship  is  evident.  The  relations  are  not  absolute 
in  every  instance,  but  there  can  be  no  doubt  whatever  of  the  tendency 
toward  decreased  absorption  as  growth  increases,  or  that  antagonism 
between  ions  results  in  decreased  absorption  of  at  least  some  of  the 
ions  present  in  the  nutrient  solution. 

We  turn  now  to  a  consideration  of  the  effects  of  a  few  of  the  salts 
of  the  heavy  metals  upon  growth  and  absorption.  In  table  10  the 
effects  of  adding  various  concentrations  of  aluminum  chloride  are 
shown.  Growth  is  decreased  in  every  concentration  of  the  salt  used. 
The  high  percentage  of  magnesium  is  marked  in  both  roots  and  tops. 
On  the  other  hand,  the  percentage  of  calcium  is  increased  relatively 
little.  The  percentage  of  iron  found  was  practically  constant  and  in 
total  quantity  is  in  marked  contrast  to  the  last  series  considered  in 
which  the  amount  was  so  small  that  it  could  not  be  determined. 

In  a  solution  of  .20  M.  calcium  chloride,  the  results  with  the  vary- 


191S]  Waynick:    Antagonism  and  Cell  Permeability  149 

ing  concentrations  of  aluminum  chloride  are  shown  in  table  11.  In 
general  the  toxic  effects  of  the  two  salts  seem  to  be  additive,  that  is. 
the  growth  in  this  series  in  which  two  salts  are  present  together  is 
less  than  in  the  preceding  series  where  aluminum  chloride  was  used 
alone.  The  decrease  is  not  great  from  the  standpoint  of  total  weight, 
but  proportionately  is  very  considerable,  amounting  to  from  33  per 
cent  to  100  per  cent  in  the  various  concentrations  employed.  The 
percentage  of  magnesium  in  the  two  series  is  about  the  same.  The 
amount  of  calcium  absorbed,  on  the  other  hand,  is  increased  over  300 
per  cent  and  remains  constant  throughout.  The  total  absorption  with 
respect  to  calcium  and  magnesium,  at  least,  is  uniformly  high.  This 
fact  is  reflected  in  the  increase  in  the  percentage  of  ash  over  that  of 
the  control.  In  the  next  series  all  factors  are  the  same  except  that 
magnesium  chloride  was  used  instead  of  calcium  chloride,  there  being 
no  difference  whatever  in  partial  or  total  concentration.  The  antag- 
onism shown  between  magnesium  chloride  and  aluminum  chloride  in 
culture  4  is  very  marked,  and  correlated  with  the  increased  growth  is 
the  marked  decrease  in  the  percentage  of  both  magnesium  and  cal- 
cium found  in  tops  and  roots.  The  percentage  of  magnesium  found 
in  the  plants  is  not  proportional  to  the  concentration  in  the  solution 
as  was  true  with  calcium  chloride.  An  interesting  case  of  the  in- 
creased absorption  of  one  element  with  a  decrease  in  the  other  is  well 
illustrated  in  the  case  of  culture  6  of  this  series.  Such  a  relationship 
has  been  noted  previously,  but  is  apparently  of  no  direct  importance 
from  the  standpoint  of  growth. 

Ferric  chloride,  a  second  trivalent  salt,  was  used  in  the  nutrient 
solution  in  the  concentration  shown  in  table  13.  In  the  concentration 
employed,  growth  is  nearly  normal  and  absorption  is  very  nearly  the 
same  as  Avith  plants  in  the  control  cultures,  except  in  the  case  of  cal- 
cium. The  decrease  in  some  instances  in  the  percentage  of  calcium 
found,  as  iron  increases  in  the  nutrient  solution,  is  notable,  and  will 
be  referred  to  later  in  connection  with  the  action  of  ferric  and  zinc 
sulphates. 

The  effects  of  adding  .20  M.  calcium  chloride,  together  with  vari- 
ous concentrations  of  ferric  chloride,  are  given  in  table  14.  The 
growth  of  roots  and  top  parallel  each  other  closely.  Marked  toxic 
effects  are  evident  in  certain  combinations  as  in  cultures  3  and  7. 
The  percentage  of  calcium  found  in  both  roots  and  tops  is  high  in 
plants  grown  in  the  same  cultures.  The  magnesium  present  in 
the  tops  shows  the  same  relationships  as  the  calcium,  although  the 


150  University  of  California  Publications  in  Agricultural  Sciences         [Vol.3 

amount  absorbed  varies  but  little  from  that  of  the  control.  In  the 
roots  magnesium  is  present  in  large  amount  when  growth  is  low  in 
culture  2,  but  in  succeeding  cultures  the  percentage  found  falls  off 
sharply  and  remains  abnormally  low  without  any  relation  to  growth 
or  concentration  of  the  solution.  The  percentage  of  iron  is  high  in 
cultures  6  and  7,  in  which  the  weight  of  the  plants  was  small. 

Substituting  magnesium  chloride  in  equivalent  concentration  for 
the  calcium  chloride  used  in  the  preceding  series,  the  results  are  of  a 
very  different  order  from  those  in  table  15.  The  absolute  growth  of 
the  tops  is  greater  than  in  series  14.  Root  growth  does  not  parallel 
the  growth  of  the  tops.  The  toxicity  of  the  solution  is  scarcely  evident 
at  some  concentrations  while  markedly  increased  at  others.  Absorp- 
tion, with  the  exception  of  the  magnesium  in  the  roots,  is  usually  low, 
amounting  to  about  that  of  the  control,  but  the  percentages  of  calcium 
and  magnesium  found  bear  no  apparent  relation  to  the  differences  in 
growth.  Iron,  however,  shows  the  inverse  relation  already  noted  in 
many  other  series  with  calcium  and  magnesium,  that  is,  high  percent- 
age present  when  growth  is  low,  and  vice  versa.  The  toxic  and  antag- 
onistic effects  as  well  may  be  due  in  this  instance  to  the  ferric  ion,  but 
this  statement  is  by  no  means  indisputable. 

In  several  tables  following,  the  effects  of  copper  salts  are  given. 
Previously  copper  salts  have  been  shown  to  be  highly  toxic  to  plants 
as  well  as  to  a  wide  variety  of  vegetative  forms.  That  they  may  also 
be  stimulating  has  been  shown  recently  by  Forbes28  using  solution 
cultures,  and  by  Lipman  and  Gericke29  in  soil  cultures.  The  reader  is 
referred  to  the  latter  paper  for  an  extensive  review  of  the  subject. 

The  results  with  copper  chloride  are  reported  in  table  16.  Growth, 
especially  that  of  the  roots,  was  limited  in  every  concentration  re- 
ported. In  fact,  the  growth  of  the  roots  was  so  limited  that  their 
weights  are  not  given.  There  is  a  suggestion  of  antagonistic  action 
between  the  nutrient  solution  and  copper  chloride  in  cultures  3  and  5. 
The  percentage  of  magnesium  found  is  high  where  growth  is  low. 
The  same  is  not  true  of  calcium,  the  percentage  of  which  is  low  and 
decreases  as  growth  decreases  to  a  certain  extent.  A  trace  of  copper 
was  found  in  every  case  and  appreciable  amounts  had  penetrated 
the  plant  tissue  at  the  two  higher  concentrations.  When  ferric 
chloride  is  added  together  with  copper  chloride  marked  antagonism  is 
shown.     Table  17  will  make  this  effect  evident.     In  this  series,  as  in 


zs  Univ.  Calif.  Publ.  Agr.  Sci.,  vol.  1   (11)17),  p.  395. 
20  Ibid.,  p.  495. 


191S]  Waynick:    Antagonism  and  Cell  Permeability  151 

several  following,  the  concentrations  of  both  salts  added  increase,  that 
is,  both  increasing  but  bearing  the  same  ratio  between  the  two. 
There  is  an  increase  of  approximately  100  per  cent  in  the  dry  weight 
of  culture  2  over  cultures  1  and  3.  The  low  absorption  of  culture  2 
as  related  to  1  and  3  is  evident.  There  is  a  marked  decrease  in  the 
percentages  of  calcium  and  magnesium  found  in  the  plants  grown  in 
culture  5,  in  which  the  dry  weight  of  the  plants  was  also  low.  At  this 
second  point,  however,  iron  and  copper  were  found  in  larger  amounts 
than  at  any  other  concentration  used.  As  in  the  previous  series  the 
percentage  of  calcium  in  the  tops  does  not  seem  to  parallel  that  in  the 
roots  or  of  magnesium  in  either  roots  or  tops.  A  similar  relationship 
was  brought  out  in  the  previous  series  in  which  copper  chloride  alone 
was  used.  No  apparent  precipitation  took  place  upon  the  addition  of 
iron  in  the  concentrations  given,  but  a  precipitate  composed  of  ferric 
phosphate  was  present  at  the  time  of  harvesting.  It  is  possible  that 
double  salts  of  copper  or  iron  with  calcium  or  magnesium  and,  for 
instance,  the  phosphate  ion  were  formed  at  the  higher  concentrations. 
Their  complexes  may  not  be  taken  up  by  the  plants  and  hence  actual 
starvation  as  far  as  these  elements  are  concerned,  may  be  responsible 
for  the  low  amounts  found  in  the  plants.  Such  a  condition  contrasts 
directly  with  one  in  which  there  is  low  permeability  due  to  antagonistic 
effects  between  the  ions  in  the  solution. 

In  table  18  mercuric  chloride  was  used  with  copper  chloride,  since 
it  was  desired  to  determine  the  effects  produced  by  the  addition  of 
two  highly  toxic  salts  to  the  nutrient  solution.  The  results  with  mer- 
curic chloride  alone  are  given  in  table  26.  They  are  somewhat  irregu- 
lar, but  there  can  be  no  doubt  of  the  correlation  between  the  quanti- 
tative presence  of  calcium  and  magnesium  in  the  tops,  of  magnesium 
in  the  roots,  and  growth.  There  is  evidence  of  a  distinct  antagonistic 
action  between  copper  and  mercuric  chlorides  both  from  the  stand- 
point of  growth  and  that  of  absorption.  The  root  growth  was  very 
limited.  The  percentage  of  calcium  and  magnesium  in  the  roots  was 
very  high ;  high  enough  to  account  for  the  decreased  growth  by  itself 
if  we  use  the  results  of  other  series  in  interpreting  this  one.  Not 
enough  iron  was  present  in  any  culture  to  permit  of  its  determination. 

Considering  the  most  common  salt  of  copper  used  in  solution  cul- 
tures and  soil  work,  the  results  as  given  in  table  19  are  especially 
noteworthy.  The  concentrations  of  the  sulphate  used  are  low.  Dis- 
tinct evidence  of  the  toxic  effects  of  the  salt,  together  with  only  slight 
decrease  in  growth  in  culture  4  of  the  series  is  shown.    High  percent- 


152  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

ages  of  calcium  and  magnesium  accompany  low  growth;  low  percent- 
ages of  calcium  and  magnesium  go  with  much  increased  growth.  No 
iron  could  be  quantitatively  determined  in  cultures  8  and  9.  The 
copper  content  shows  no  variations  which  may  be  regarded  as  impor- 
tant, in  fact  the  amount  taken  up  by  the  plants  is  somewhat  lower 
where  decreased  growth  is  shown. 

Zinc  sulphate  was  used  with  copper  sulphate  as  shown  in  table  20. 
There  is  little  evidence  of  antagonism  between  the  two  salts.  At  the 
same  time  there  is  evidently  no  direct  relationship  between  concen- 
tration and  toxic  effect,  since  growth  does  not  decrease  regularly  with 
increasing  concentration.  While  the  percentages  of  calcium  and  mag- 
nesium found  are  somewhat  irregular,  they  increase  rapidly  as  growth 
becomes  less.  The  percentage  of  magnesium  found  in  the  tops  in 
culture  8  was  1.10  per  cent,  and  in  the  roots  1.91  per  cent.  This 
occurred  with  the  same  concentration  of  the  magnesium  ion  in  the 
nutrient  as  in  culture  1.  The  percentage  of  copper  found  in  the  dry 
matter  is  distinctly  larger  than  that  found  in  the  preceding  series,  in 
which  copper  sulphate  alone  was  used. 

Copper  sulphate  used  with  ferric  sulphate  shows  no  evidence  of 
antagonism  between  the  two  if  the  growth  of  the  tops  alone  is  con- 
sidered, but  with  the  roots  there  is  a  marked  increase  in  growth  in 
cultures  3  and  4  of  the  series.  The  percentages  of  magnesium  found 
in  the  roots  is  low  and  constant,  which  contrasts  markedly  with  the 
amounts  determined  in  the  previous  series.  The  calcium  likewise 
varies  but  little  in  the  tops  and  its  percentage  remains  low.  On  the 
other  hand,  the  percentages  of  calcium  in  the  tops  and  magnesium  in 
the  roots  show  marked  increases  as  growth  decreases.  The  amount 
of  iron  remains  very  uniform  until  the  last  culture  of  the  series  is 
reached,  when  a  marked  increase  is  recorded.  It  will  be  noted  that  the 
percentage  of  calcium  decreases  to  nearly  one-third  of  the  original  in 
the  same  culture.  This  relation  has  been  noted  previously  in  other 
series. 

The  stimulation  resulting  from  the  addition  of  ferric  sulphate  to 
the  nutrient  solution  in  the  concentrations  given  in  table  22  is  remark- 
able, a  total  dry  weight  of  3.9016  grams  for  the  tops  of  six  plants 
being  recorded.  The  growth  of  the  roots  does  not  parallel  that  of 
the  tops.  In  the  highest  concentration  of  ferric  sulphate  employed, 
the  root  growth  decreased  while  the  growth  of  the  tops  was  increased. 
Attention  has  already  been  called  to  cases  of  this  kind  in  which 
there  may  be  an   increase  in  the  growth  of  tops  with  a  decrease  in 


191S]  Waynick:    Antagonism  and  Cell  Permeability  153 

the  root  growth,  or  vice  versa.  As  will  be  noted,  the  percentages 
of  calcium  and  magnesium  found  are  low,  in  fact  below  the  control 
in  every  case.  Whether  or  not  ferric  sulphate  would  be  stimulating 
in  still  higher  concentrations  is  not  known,  but  it  is  probable  that  the 
limit  of  stimulation  was  reached,  since  the  roots  show  a  marked  de- 
crease in  growth  in  the  highest  concentration  used.  The  percentage 
of  iron  found  is  comparatively  high.  The  reason  for  this  increased 
growth  is  evidently  bound  up  with  the  presence  of  the  ferric  salt, 
but  no  idea  of  the  nature  of  its  action  can  be  given.  It  is  very  evi- 
dent from  the  present  data,  however,  that  the  amounts  of  the  elements 
present  in  the  plants  were  low. 

In  table  23  the  results  with  zinc  sulphate  alone  are  reported. 
There  is  no  stimulation  or  no  antagonism  between  zinc  sulphate  and 
the  other  constituents  of  the  solution  evident  in  any  concentration. 
As  growth  decreases  magnesium  was  found  present  in  larger  amounts 
than  in  the  cultures  in  which  growth  was  more  nearly  normal.  The 
percentage  of  calcium  remains  very  much  the  same  in  the  tops  and 
decreases  rapidly  in  the  roots  with  decreasing  growth.  Here  we  have 
a  suggestion  of  a  relationship  between  zinc  and  calcium  as  has  already 
been  referred  to  in  the  case  of  iron.  It  can  only  be  stated,  however, 
that  the  results  as  regards  calcium  penetration  are  exceptional  in  the 
light  of  the  results  in  other  series  previously  referred  to. 

Turning  to  table  24,  in  which  the  results  with  zinc  sulphate  and 
ferric  sulphate  are  given,  there  is  a  marked  contrast  on  the  one  hand 
with  series  20  in  which  zinc  sulphate  and  copper  sulphate  were  used, 
and  on  the  other  hand  with  the  preceding  series  in  which  zinc  sulphate 
alone  was  used.  In  this  series  there  is  marked  antagonism  shown  be- 
tween the  salts  employed.  This  is  true  for  both  tops  and  roots,  but 
the  most  marked  increase  in  both  does  not  occur  in  the  same  culture. 
The  marked  increase  in  growth  of  the  tops  evident  in  culture  4  is 
accompanied  by  a  decrease  in  the  percentages  of  calcium  and  mag- 
nesium present  in  the  tops  but  not  in  the  roots.  The  percentage  of 
magnesium  in  the  roots  increases  with  decreased  growth  throughout 
the  series.  The  calcium  in  the  tops  is  low  and  abnormally  so  in  the 
roots.  Growth  is  good  throughout  the  series  and  in  culture  4  is  in- 
creased about  50  per  cent  above  the  control.  This  result  would  hardly 
be  expected  from  the  decreases  recorded  where  zinc  sulphate  was  used 
alone  in  the  preceding  series.  The  percentage  of  iron  varies  some- 
what, but  does  not  increase  or  decrease  with  any  regularity  in  any 
one  direction.  Attention  is  again  called  to  the  low  calcium  content, 
especially  of  the  roots. 


154  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

Little  can  be  said  of  the  mercuric  chloride  ferric  sulphate  series 
given  in  table  25.  Growth  is  uniformly  low  throughout,  with  con- 
siderable variation  between  duplicate  cultures.  The  percentage  of 
magnesium  is  very  high  in  the  roots  and  while  less  in  the  tops,  is 
much  above  that  of  the  control.  The  percentage  of  calcium  is  uni- 
formly low  in  both  tops  and  roots.  Attention  is  called  to  the  fact 
that  no  iron  could  be  determined  quantitatively,  except  in  the  highest 
concentration  of  salts  used.  This  condition  is  striking  when  the 
rather  large  amounts  of  ferric  sulphate  in  the  solution  are  considered. 

A  short  series  is  reported  in  table  26  in  which  the  toxic  effects 
of  mercuric  chloride  when  used  alone,  are  evident.  There  is  a  de- 
crease in  growth  with  increasing  concentration  of  the  added  salt  and 
also  an  increasing  percentage  of  both  calcium  and  magnesium  found. 
The  very  low  ash  content  given  by  the  plants  in  this  series  is  of 
interest  and  will  be  discussed  below. 


External  Appearances  of  the  Plants 

It  seems  worth  while  to  note  here  a  few  of  the  more  striking 
appearances  of  the  plants.  Since  iron  salts  were  purposely  excluded 
from  all  solutions  except  those  in  which  it  was  planned  to  study  their 
effects,  the  control  plants  were  of  a  more  or  less  yellowish  green  color. 
Aside  from  this  no  differences  were  noted  between  control  plants 
grown  with  or  without  the  addition  of  a  little  ferric  phosphate  to 
the  nutrient. 

In  every  series  in  which  growth  was  limited  by  the  presence  of 
magnesium  salts  the  roots  were  short  and  much  thickened.  With  a 
high  concentration  of  magnesium  in  a  balanced  solution,  this  effect 
was  not  noted  however.  High  concentrations  of  magnesium  were  also 
apparent  from  the  decided  yellowing  of  the  older  leaves.  Excessive 
amounts  of  calcium  were  characterized  by  the  appearance  of  brown 
spots  or  streaks  on  the  leaves.30 

When  any  considerable  growth  was  permitted  the  plants  grown  in 
solutions  of  copper  salts  were  dark  green  in  color.31  Where  growth 
was  good  the  roots  were  apparently  normal.  In  several  of  the  higher 
concentrations  used,  copper  hydroxide  was  deposited  upon  the  roots, 
especially  about  the  tips.  A  suggestion  is  made  that  possibly  copper 
may  replace  iron  as  a  catalyzer  in  connection  with  the  building  or 
activation  of  chlorophyll. 


30  Jost,  Plant  physiology  (Oxford,  Clarendon  Press,  1907),  p.  85. 
f"  CJniv.  Calif.  Agr.  Sci.,  vol.  1    (1917),  pp.  495-588. 


1918]  Waynick:    Antagonism  and  Cell  Permeability  155 

Several  cultures  in  which  mercuric  chloride  was  used  and  in  which 
growth  was  good,  displayed  the  same  dark  green  color  as  noted  for 
copper  salts  and  the  same  suggestion  as  made  for  the  functioning  of 
copper  in  this  color  relationship  may  hold  for  mercuric  salts  as  well 
in  very  dilute  solutions. 

The  color  was  light  green  when  iron  salts  were  present;  with  the 
other  salts  used  no  marked  external  effects  were  noted. 

General  Review  of  Experimental  Results 

It  seems  advisable  to  consider  the  results  reported  in  the  previous 
tables  together,  so  that  the  data  presented  in  one  table  may  be  more 
closely  correlated  with  those  given  in  another.  It  is  proposed  to  do 
this  in  the  present  section  and  further  to  discuss  briefly  the  more 
important  relationships  shown. 

It  will  be  noted  in  the  accompanying  tables  that  there  is  consid- 
erable variation  between  the  controls  grown  at  different  seasons  of 
the  year.  This  was  to  be  expected,  since  conditions  in  the  green- 
house varied  between  the  different  growing  periods.  For  this  reason 
it  is  not  possible  to  compare  one  series  of  cultures  with  another  so 
far  as  absolute  weights  of  the  dry  matter  are  concerned.  Within  any 
one  series  or  between  series  grown  at  the  same  time  the  absolute 
weights  are  comparable.  This  point  must  be  borne  in  mind  in  con- 
sidering the  results  as  a  whole.  In  some  cultures,  however,  growth 
was  stimulated  to  such  an  extent  as  to  far  surpass  any  variation 
between  series  due  to  differing  external  conditions.  Such  a  case  is 
that  of  series  22,  in  which  ferric  sulphate  was  added  to  the  nutrient 
solution  in  varying  amounts.  In  culture  5  of  this  series,  the  dry 
weight  was  over  twice  that  of  any  control  plants  grown  during  the 
entire  time. 

The  experimental  work  with  the  salts  of  calcium  plus  magnesium 
was  rather  extensive.  McCooP2  has  reviewed  the  previous  work  with 
calcium  and  magnesium  salts  as  related  to  plants,  so  a  discussion  of 
that  phase  of  the  relationships  between  the  two  need  not  be  entered 
into  here.  In  his  own  work  McCool  found  that  calcium  chloride  was 
effective  in  antagonizing  the  poisonous  effects  of  magnesium  chloride 
and  magnesium  sulphate.  He  found  a  slight  increase  in  the  growth 
of  pea  seedlings  over  the  controls  based  upon  the  green  weight  of  the 
plants.  This  was  the  case  in  distilled  wTater  and  in  nutrient  solution. 
It  seems  probable  that  the  nutrient  solution  used  by  McCool  was  not 


32  Cornell  Univ.  Agr.  Exp.  Sta.  Mem.  2   (1913),  p.  129. 


156  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

a  balanced  solution,  since  the  addition  of  either  magnesium  or  cal- 
cium chloride  resulted  in  an  increased  growth  of  the  pea  seedlings. 

In  the  present  investigation  there  are  only  two  cases  in  which  the 
growth  of  the  plants  was  greater  with  both  calcium  and  magnesium 
chlorides  present  than  when  calcium  chloride  was  used  alone  in  vari- 
ous concentrations,  one  in  culture  6,  series  2,  the  other  in  culture  11, 
series  3.  In  the  latter  culture  the  dry  weight  of  the  plants  was  twice 
that  in  the  same  concentration  of  calcium  chloride  alone.  There  are 
marked  differences  in  growth  recorded  between  different  combinations 
and  concentrations  of  the  two  salts,  and  as  can  be  easily  seen  from 
the  graphs,  the  percentages  of  the  two  ions  found  in  the  plants  show 
an  inverse  relation  to  growth  in  nearly  every  instance.  Proceeding 
from  series  to  series,  the  amount  of  magnesium  found  in  the  plants 
increases  with  the  concentration  of  the  magnesium  chloride  in  the 
nutrient  solution. 

Magnesium  sulphate  is  not  as  toxic  as  magnesium  chloride  in 
equivalent  concentrations  of  the  kation.  Growth  in  solutions  of  mag- 
nesium sulphate  plus  calcium  chloride  was  superior  in  every  case  to 
that  found  when  the  salts  were  used  separately.  There  is  a  marked 
contrast  between  calcium  chloride  and  calcium  nitrate  in  antagoniz- 
ing the  toxic  effects  of  magnesium  sulphate,  the  nitrate  proving  less 
effective  than  the  chloride  in  concentrations  of  .12  M.  and  over.  This 
is  of  especial  interest,  since  the  qualitative  ionic  relations  of  the  nutri- 
ent are  not  altered.  It  is  possible  that  we  are  dealing  with  the  effects 
of  undissociated  molecules  in  the  higher  concentrations,  which  may 
be  very  different  from  ionic  effects. 

Results  with  Salts  of  the  Heavy  Metals 

Since  salts  of  aluminum,  copper,  zinc,  iron  and  mercury  were 
used,  it  will  be  necessary  for  the  sake  of  clearness  to  treat  each  more 
or  less  separately. 

Miyake33  has  shown  aluminum  chloride  to  be  highly  toxic,  in  con- 

N 
centrations  above     — -,  to  rice  seedlings   grown  in  water  cultures. 

Similar  results  have  been  reported  by  House34  and  Gies,  Micheels  and 
De  Heen,35  Duggar,36  and  Ruprecht,37  working  with  several  aluminum 


33  Jour.  Biol.  Chem.,  vol.  25   (1916),  p.  23. 
84Amer.  Jour.  Physiol.,  vol.  15   (1905),  p.  19. 

35  Bull.  Acad.  Roy.  Belg.   (1905),  p.  520. 

36  Plant  Physiology,  New  York,  Macmillan,  1911, 

37  Mass.  Exp.  Sta.  Bull.  161   (1915),  p.  125. 


1918]  Waynick:    Antagonism  and  Cell  Permeability  157 

salts.  Probably  the  work  of  Abbott,  Conner  and  Smalley38  is  of  more 
direct  interest  here.  These  investigators  found  aluminum  nitrate  to 
be  toxic  to  corn  seedlings  in  the  presence  of  nutrient  solutions. 
E.  Kratzmann39  has  reported  stimulation  due  to  the  presence  of  small 
amounts  of  aluminum  salts.  Miyake40  concludes  further  that  the 
effects  observed  with  aluminum  chloride  cannot  be  attributed  to  the 
hydrogen  ion  resulting  from  the  dissociation  of  the  salt. 

Aluminum  chloride  was  found  to  be  toxic  in  every  concentration 
used  in  the  present  work.  The  effect  of  the  presence  of  calcium 
chloride  in  a  concentration  of  .20  M.  was  to  decrease  growth  still  fur- 
ther, indicating  that  its  toxic  effect,  as  reflected  in  growth,  was  but 
additive  to  that  of  aluminum  chloride.  With  magnesium  chloride 
present  in  equivalent  concentration  as  the  calcium  chloride,  there  is  a 
marked  antagonism  at  a  concentration  of  .000066  M.  of  aluminum 
chloride  with  .20  M.  magnesium  chloride.  The  increase  in  dry  weight 
was  100  per  cent  greater  than  in  an  equivalent  concentration  of 
aluminum  chloride  alone  and  300  per  cent  greater  than,  with  mag- 
nesium chloride  in  the  concentration  given.  This  culture  has  been 
referred  to  especially  since  it  furnishes  a  striking  example  of  antag- 
onism between  bivalent  and  trivalent  salts,  both  of  which  are  highly 
toxic  when  used  alone.  The  chloride  ion  was  a  constant  as  far  as  this 
and  the  preceding  series  are  concerned,  the  only  difference  between 
the  two  cases  being  the  use  of  calcium  chloride  in  one  and  magnesium 
chloride  in  the  other.  It  seems  logical  to  conclude  that  the  action  is 
specific  as  regards  the  magnesium  and  aluminum  ions.  Whatever 
the  nature  of  this  action  may  be,  it  is  certainly  not  shown  between 
calcium  and  aluminum  ions. 

The  same  general  relationships  are  brought  out  between  ferric 
chloride  and  calcium  and  aluminum  chlorides.  Ferric  chloride  did 
not  prove  toxic  in  the  concentrations  used,  growth  differing  but  little 
from  that  of  the  control.  When  calcium  chloride  was  present  in  a 
concentration  of  .20  M.  throughout  the  series,  growth  was  half  or  less 
than  half  that  recorded  when  ferric  chloride  alone  was  present. 
Magnesium  chloride  in  equivalent  concentrations,  as  the  calcium 
chloride  above,  affected  growth  but  little.  In  other  words,  magnesium 
chloride  did  not  prove  toxic  in  the  presence  of  certain  concentrations 
of  ferric  chloride.  The  relations  between  the  four  salts  may  be  briefly 
summarized  as  follows :   There  is  no  antagonism  shown  between  alumi- 


38  Incl.  Exp   Sta.  Bull.  170  (1913),  p.  329. 

39Chem.  Ztg.,  vol.  38  (1914),  p.  1040. 

40  Jour.  Biol.  Chem.,  vol.  25  (1916),  p.  23. 


158  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

num  chloride  and  calcium  chloride.  There  is  very  little,  if  any,  be- 
tween ferric  chloride  and  calcium  chloride.  Magnesium  chloride  and 
ferric  chloride  show  marked  antagonism  in  all  concentrations  used 
as  do  magnesium  chloride  and  aluminum  chloride  in  certain  concen- 
trations of  the  two  salts.  Magnesium  chloride  and  ferric  chloride 
show  marked  antagonism  in  all  concentrations  as  do  magnesium 
chloride  and  aluminum  chloride  in  one  concentration  of  the  latter  salt. 

Reference  has  already  been  made  to  Miss  Brenchley's  monograph41 
and  to  the  paper  by  Lipman  and  Gericke,42  in  which  the  literature 
relating  to  the  effects  of  copper,  zinc,  and  iron  salts  on  plants  is 
reviewed.  Suffice  it  to  say  that  the  results  reported  by  different 
investigators  are  very  conflicting,  due  largely  to  the  widely  different 
methods  used  and  the  varying  conditions  under  which  the  various 
data  were  obtained. 

In  the  present  work,  copper  chloride  was  toxic  in  every  concen- 
tration used.  There  was  marked  antagonism  between  copper  and 
ferric  chlorides  both  from  the  standpoint  of  growth  and  of  absorption. 

Copper  sulphate  did  not  prove  to  be  uniformly  toxic.  Growth  was 
nearly  normal  in  one  concentration  used  while  very  much  diminished 
in  a  lower  concentration.  The  term  stimulation  might  be  applied 
here,  but  in  the  present  discussion  it  is  applied  only  when  growth  due 
to  the  presence  of  an  added  salt  or  salts  is  undoubtedly  greater  than 
that  in  the  control. 

Toxic  effects  are  correlated  with  increased  absorption  and  antag- 
onistic effects  with  decreased  absorption  as  in  other  series  reported. 

Growth  was  always  less  with  zinc  sulphate  present  in  the  nutrient 
solution  than  in  the  latter  alone.  Copper  and  zinc  sulphate  together 
were  no  more  toxic  than  a  solution  of  zinc  sulphate  alone. 

The  case  with  ferric  sulphate  is  clearly  one  of  stimulation.  The 
dry  weight  was  over  twice  that  of  the  controls  in  one  concentration 
of  the  salt  used  and  far  superior  in  several  concentrations  to  that 
of  the  plants  grown  in  the  controls.  Wolff43  has  reported  similar 
results  when  iron  was  used  in  the  form  of  the  citrate,  an  increase 
in  growth  comparable  to  that  noted  above  having  been  obtained.  He 
found  further  that  nickel  or  chromium  could  not  be  used  to  replace 
iron. 

The  toxic  effects  of  copper  sulphate  were  markedly  reduced  by 
the  presence  of  ferric  sulphate  when  we  consider  the  results  as  a 


11  Inorganic  plant  poisons  and  stimulants.     1915. 
'-  Univ.  Cal.  Pub.  Agr.  Sci.,  vol.  1   (1917),  p.  395. 
43C.-E.  Acad.  Sci.  (Paris),  vol.  157   (1913),  p.  1022. 


1918]  Waynicl*:    Antagonism  and  Cell  Permeability  159 

whole,  although  in  one  instance  growth  was  greater  with  copper  sul- 
phate alone  than  when  both  salts  were  added  together. 

The  second  case  of  stimulation  was  noted  with  zinc  sulphate  and 
ferric  sulphate  in  certain  concentrations.  In  series  26  four  cultures 
gave  growth  superior  to  that  obtained  in  the  control  for  the  series, 
and  throughout  growth  was  good  when  the  two  salts  referred  to  above 
were  present  together,  over  the  range  of  concentrations  employed. 
Low  absorption  was  noted.  In  summarizing  the  relations  of  ferric, 
cupric  and  zinc  sulphates,  it  is  evident,  from  the  discussion  above, 
that  zinc  sulphate  was  toxic  in  every  concentration  used.  Copper  sul- 
phate was  toxic,  but  marked  variation  in  degree  was  shown  between 
various  concentrations.  Ferric  sulphate  was  stimulating.  Copper 
sulphate  and  zinc  sulphate  were  no  more  toxic  together  than  when 
each  was  used  alone.  Ferric  sulphate  modified  somewhat  the  toxic 
effects  of  copper  sulphate.  Zinc  sulphate  and  ferric  sulphate  together 
proved  stimulating  to  the  growth  of  plants.  As  contrasted  with  the 
chlorides,  the  sulphates  of  copper  and  iron  were  less  toxic  to  barley 
over  the  range  of  concentrations  used  in  this  investigation. 

Taking  the  results  as  a  whole,  twelve  instances  of  a  marked  in- 
crease in  growth  at  certain  definite  concentrations  of  one  or  more 
added  salts  have  been  noted.  With  every  such  increase  there  is  a  very 
notable  decrease  in  the  amount  of  calcium  and  magnesium  absorbed. 
The  increase  in  growth  is  attributed  to  antagonistic  salt  action;  de- 
creased absorption  is  undoubtedly  due  to  the  same  action,  which 
tends  to  preserve  the  normal  permeability  of  the  plasma  membrane. 

In  addition  to  the  twelve  instances  referred  to  above,  we  find 
in  series  after  series,  the  toxic  effects  of  the  solution  in  which  the 
plants  were  growing,  noticeable  not  alone  by  decreased  growth  but 
also  by  increased  absorption.  The  roots  and  tops  may  not  show  the 
same  relations  as  regards  the  amounts  of  calcium  and  magnesium 
taken  up.  For  example,  in  series  25,  in  which  ferric  sulphate  and 
mercuric  chloride  were  used  together,  the  toxicity  of  the  solutions  was 
evident  by  the  very  limited  growth,  yet  the  composition  of  the  tops 
was  about  normal.  In  the  roots,  however,  the  percentage  of  mag- 
nesium was  found  to  be  tremendously  increased. 

It  is  of  interest  to  refer  again  to  the  very  low  ash  content  and 
relatively  low  absorption,  considering  the  very  limited  growth,  in  the 
few  cultures  in  which  mercuric  chloride  was  used  alone.  It  is  pos- 
sible that  relatively  large  amounts  of  mercuric  salts  were  taken  up 
by  the  plants  which  were  volatilized  on  ashing  the  residue ;  thus  the 
low  percentage  of  ash  may  be  less  surprising. 


160  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 

Possible  Effects  of  Variations  in  the  Concentrations  of 
the  Solutions  on  the  Plants 

No  attempt  was  made  to  maintain  the  total  concentration  of  the 
nutrient  solution  constant.  This  would  be  exceedingly  difficult  to  do 
in  work  of  this  character,  since  it  would  be  necessary  to  vary  the 
concentration  of  the  nutrient  solution  to  maintain  the  balance  of  the 
solution  as  regards  total  concentration.  The  conclusion  seems  justi- 
fied that  within  the  range  employed  the  concentration  of  the  nutrient 
solution  is  of  minor  importance  as  far  as  growth  is  concerned.  For 
instance,  in  table  1,  the  variation  in  the  concentration  of  the  solution 
was  .279  M.  in  terms  of  calcium  chloride,  yet  the  total  growth  varied 
but  little  from  .001  M.  to  .28  M.  Again  in  table  2  the  growth  is  very 
nearly  the  same  at  a  concentration  of  .25  M..  with  calcium  and  mag- 
nesium chlorides,  and  a  total  concentration  of  .54  M.  of  the  same  salts. 
In  table  3  the  greatest  growth  occurred  in  a  concentration  of  .46  M. 
in  terms  of  the  salts  above  mentioned,  while  at  the  lower  concentra- 
tions of  .304  M.,  growth  was  but  a  third  that  obtained  in  the  higher 
concentrations.  These  examples  make  clear  the  point  above  men- 
tioned, namely,  that  the  concentration  over  the  range  used  was  of 
but  minor  importance.  It  is  obvious  that  the  above  discussion  does 
not  apply  to  the  series  in  which  salts  of  the  heavy  metals  were  used, 
since  the  variations  in  concentration  in  those  series  were  but  slight. 

Consideration  of  a  Possible  Calcium-Magnesium  Ratio 

Since  Loew44  first  advanced  the  hypothesis  of  the  lime-magnesia 
ratio,  much  experimental  evidence  has  been  collected  by  various  inves- 
tigators both  for  and  against  the  existence  of  an  optimum  ratio  be- 
tween these  two  elements  as  regards  the  growth  of  plants.  The 
literature  bearing  upon  the  subject  has  been  very  fully  reviewed  by 
Lipman,45  so  that  detailed  references  are  not  necessary  here. 

Since  the  ratios  of  calcium  to  magnesium  in  the  solution  used  by 
the  writer  were  known  and  also  because  of  the  fact  that  the  analytical 
data  allowed  of  the  calculation  of  such  a  ratio  for  the  plants,  it 
seemed  of  interest  to  present  some  of  these  data  here. 

The  following  two  tables  give  the  results  obtained  from  two  series 
in  which  widely  varying  proportions  of  calcium  and  magnesium  were 
used. 


44  Flora,  vol.  75    (1892),  p.  368. 

>■>  Plant  world,  vol.  19   (1916),  p.  83. 


1918] 


Waynick:    Antagonism  and  Cell  Permeability 


161 


Table 

,  27 

Rati 
Mgto 
in  soli, 

41 

0 

Ca 
tion 

1 

Dry  weight 
tops 

.3536 

Ratio 

Mg  to  Ca 

in  tops 

2.3   :   1 

Dry  weight, 
roots 

.1218 

Ratio 
Mg  to  Ca 
in  roots 

1.2    :    1 

16 

1 

.5484 

4.6    : 

1 

.1519 

1 

2 

8 

1 

.5885 

6.0   : 

1 

.1266 

1 

4.1 

1 

.4774 

2.6   : 

1 

.1509 

3.3 

2.7 

1 

.3433 

2.6    : 

1 

.1497 

1.2 

2.0 

1 

.6775 

2.7   : 

1 

.2119 

1.3 

1.6 

1 

.4136 

1.2    : 

1 

.1500 

1.3 

1.3 

1 

.4431 

1       : 

1.2 

.1421 

1.2 

1 

1 

.3745 

1.3    : 

1 

.1138 

1 

1.3 

1 

1.2 

.3268 

1.5   : 

1 

.1254 

1 

1.5 

1 

1.4 

.2815 

1.8   : 

1 

.1053 

1 

1.2 

1 

1.8 

.5030 

1       : 

1.2 

.1044 

1.2 

Table  27  was  computed  from  the  results  given  in  table  2.  Mag- 
nesium chloride  was  present  in  uniform  concentration  of  .24  M.  with 
varying  concentrations  of  calcium  chloride. 

It  will  be  noted  that  the  dry  weights  with  a  ratio  of  magnesium  to 
calcium  of  16 :  1,  8 :  1,  and.  1 :  18  are  nearly  the  same.  The  ratios  of 
these  two  elements  found  in  the  plants  grown  in  these  solutions  were 
2  :  1,  1 :  1.  1 :  1.2  for  the  roots,  and  1 :  4.6,  1  :  1.6,  1 :  1.2  for  the  tops. 
Further,  the  dr}^  weight  of  plants  grown  in  a  solution  in  which  the 
ratio  was  41 : 1  and  with  a  ratio  of  1 :  1  are  nearly  the  same.  It  is 
evident  that  the  same  ratio  for  the  roots  may  not  hold  for  the  tops. 


Table  28 

Ratio 
Mg  to  Ca 
in  solution 

Dry  weight, 
tops 

Ratio 

Mg  to  Ca 

in  tops 

Dry  weight, 
roots 

Ratio 

Mg  to  Ca 

in  roots 

20.2    : 

1 

.3033 

5.5    :    1 

.0822 

4.4    :    1 

10.5    : 

1 

.4716 

5.4   :    1 

.1572 

4.7    :    1 

6.8   : 

1 

.2799 

4.8    :   1 

.0780 

6.3   :   1 

5.1    : 

1 

.1999 

5.8    :   1 

.0342 

6.0   :   1 

4.0   : 

1 

.1999 

4.8    :   1 

.0636 

7.4   :   1 

3.4   : 

1 

.4363 

1.3    :   1 

.1122 

4.1   :   1 

2.5   : 

1 

.4013 

1.8   :   1 

.1143 

2.8    :   1 

2.0   : 

1 

.2734 

2.3    :   1 

.0677 

4.7   :   1 

1.7   : 

1 

.2603 

2.5   :   1 

.0867 

6.8    :   1 

Table  28  gives  the  ratios  in  the  solutions  used  in  series  4,  in  which 
the  ratios  of  magnesium  to  calcium  varied  from  20.2 :  1  to  1.7 : 1. 
Growth  is  nearly  the  same  in  solutions  in  which  the  ratio  was  10.5 :  1 
as  in  those  in  which  the  ratio  is  3.4: 1  or  2.5: 1.  The  plants  grown 
in  these  cultures  gave  the  following  values  for  the  tops :  5.4 : 1,  3.4 : 1, 
2.5 :  1,  and  for  the  roots,  4.7  :  1,  4.1 :  1,  and  2.8 : 1.  There  is  a  tendency 
for  the  ratio  of  calcium  to  magnesium  in  the  plants  to  become  narrower 


162  University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 

as  the  ratio  of  these  two  ions  in  the  solution  becomes  narrower. 
Where  a  wide  ratio  exists  in  the  solution,  there  is  always  a  much  nar- 
rower ratio  in  the  plants. 

From  the  brief  discussion  above  it  is  evident  that  the  barley  plants 
grew  equally  well  in  solutions  having  widely  different  ratios  of  cal- 
cium and  magnesium  ions.  There  is  no  "optimum  lime-magnesia 
ratio,"  as  Gile46  and  Wyatt47  as  well  as  others  have  shown,  and  their 
results  are  confirmed  in  the  present  investigation. 

The  balance  between  all  the  ions  present  in  the  solution  appears 
to  be  of  far  greater  importance  than  any  single  ratio.  A  considera- 
tion of  the  ratios  existing  between  the  various  ions  of  the  nutrient 
solution,  aside  from  calcium  and  magnesium  used,  is  reserved  for 
further  study. 

Permeability  and  Antagonism 

It  is  not  proposed  to  enter  into  a  discussion  of  the  structure  and 
composition  of  the  plasma  membrane.  Davidson48  has  recently  sum- 
marized our  present  knowledge  concerning  it  with  special  reference 
to  selective  permeability.  A  discussion  of  the  various  theories  which 
have  been  advanced  to  explain  antagonistic  salt  action  need  not  be 
taken  up  in  detail  here.  The  reader  is  referred  to  papers  by  Clark,49 
Loeb,50  Osterhout,51  Loew,52  Koenig  and  Paul,53  True  and  Gies54,  True 
and  Bartlett,55  Kearney  and  Cameron,65  and  Ostwald57,  for  a  discus- 
sion of  the  various  factors  which  may  be  of  importance  in  this  con- 
nection. 

The  recent  work  of  Clowes58  and  Fenn59  is  important  and  some 
very  striking  similarities  between  the  action  of  toxic  and  antagonistic 
solutions  on  oil  emulsions  and  on  gelatine  on  the  one  hand,  and  plant 
cells  on  the  other,  have  been  reported  by  these  investigators. 


46  Porto  Eico  Exp.  Sta.,  Bull.  12   (1912). 

47  Jour.  Agr.  Research,  vol.  6  (1916),  p.  589. 

48  Plant  World,  vol.  19  (1916),  p.  331. 
40Bot.  Gaz.,  vol.  33  (1902),  p.  26. 
■r>oArchiv.  ges.  Physiol.,  vol.  88   (1902),  p.  68. 
5i  Science,  n.s.,  vol.  35   (1912),  p.  112. 

52  Flora,   vol.  75    (1892),  p.   368. 

r>3  Zeitschr.  Hygiene  u.  Infektionskranklieiten,  vol.  25  (1897),  p.  1 

54  Bull.  Torr.  Bot.  Club,  vol.  30  (1903),  p.  390. 

■>■>  U.  8.  Dept.  Agr.,  Bull.  231,  1912. 

•r'0  U.  S.  Dept.  Agr.,  Bull.  71,  1902. 

57  Archiv.  ges.  Physiol.,  vol.  120  (1907),  p.  19. 

58  Jour.  Phys.  Chem.,  vol.  20  (1916),  p.  407. 
59Proc.  Nat.  Acad.  Sci.,  vol.  2  (1916),  p.  539. 


1918]  IV  ay  nick:    Antagonism  and  Cell  Permeability  163 

To  define  normal  permeability  is  very  difficult.  There  seems  to 
be  a  comparatively  wide  range  of  concentration  of  salts  over  which 
the  amount  of  any  element  taken  up  may  vary  without  affecting  the 
growth  of  the  plant  to  any  considerable  extent.  There  is  likewise  a 
wide  range  over  which  the  ratio  of  any  one  element  to  any  other  may 
change  without  being  detrimental  to  plant  growth.  The  latter  point 
has  been  discussed  above  in  connection  with  a  possible  optimum 
calcium-magnesium  ratio  for  plants.  The  first  point  referred  to  has 
been  very  well  treated  by  Gile  and  Ageton,60  so  that  further  reference 
need  not  be  given  here. 

For  the  work  in  hand  the  percentage  composition  of  the  plants 
grown  in  the  control  cultures  seemed  to  be  the  most  logical  criterion 
of  normal  permeability  available.  There  are  variations  between  the 
controls  as  regards  composition,  but  they  are  relatively  small.  On 
the  other  hand,  the  percentages  of  magnesium,  for  instance,  range 
from  .02  per  cent  to  9.21  per  cent,  depending  upon  the  solution  used. 
The  percentages  of  calcium  differ  over  a  wide  range  as  well.  From 
the  data  presented  there  can  be  no  doubt  whatever  that  the  composi- 
tion of  the  plant,  as  regards  inorganic  constituents  at  least,  may  be 
altered  enormously  by  variations  in  the  surrounding  solution. 

That  portion  of  the  root  system  in  any  plant  which  functions  as  a 
semipermeable  membrane  is  obviously  of  greatest  importance  in  a 
study  of  the  present  kind.  The  actual  area  of  the  membrane  which 
is  in  contact  with  the  solution  must  be  known  in  every  case  before  it 
can  be  said  that  the  permeability  of  one  root  system  is  greater  than 
that  of  another.  The  actual  area  of  the  plasma  membrane  cannot  be 
measured  directly  because,  in  the  first  place,  we  have  no  means  of 
determining  just  how  much  of  the  root  is  involved,  and  secondly,  the 
area  concerned  may  be  changing  continually. 

Length  of  the  roots  and  their  number  and  length  together  as  well 
as  green  weight  and  dry  weight  have  been  taken  as  criteria  of  the 
existence  of  antagonism.  In  the  present  paper  the  dry  weight  has 
been  taken  as  proportional  to  the  area  of  the  plasma  membrane 
through  which  salts  may  enter  the  plant.  It  cannot  be  stated  defi- 
nitely that  the  two  are  proportional.  They  have  only  been  so  con- 
sidered since  the  dry  weight  of  the  plant  was  the  most  logical  criterion 
to  employ.  The  reservation  must  always  be  made  that  the  two  may 
not  be  directly  proportional,  even  though  they  are  treated  as  being  so. 

That  the  permeability  of  the  plasma  membrane  of  the  plant  cells 


so  Porto  Eico  Agr.  Exp.  Sta.  Bull.  16,  1914. 


164  University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 

is  changed  by  the  nature  and  balance  of  the  solution  surrounding  the 
roots  there  can  be  no  doubt  from  the  data  already  given.  That  a 
number  of  ions  are  capable  of  acting  in  a  very  similar  manner  to  one 
another  as  regards  permeability  is  also  evident  from  the  present  work. 
Further,  the  same  salt  may  act  differently  at  different  concentrations, 
preserving  nearly  normal  permeability  at  some  and  allowing  the  pene- 
tration of  large  numbers  of  ions  at  others.  As  previously  stated,  the 
total  balance  of  the  solution  is  of  vital  importance  in  the  preservation 
of  normal  permeability,  which  is  in  turn  correlated  with  normal 
growth. 

In  connection  with  the  salts  of  the  heavy  metals,  the  amounts  of 
the  kation  of  cupric  and  ferric  salts  which  had  penetrated  the  plant 
tissue  were  determined  in  a  number  of  instances.  The  percentages 
found  were  low.  Further,  whenever  these  salts  proved  toxic,  the 
amounts  of  calcium  and  magnesium  found  in  the  plants  were  high ; 
high  enough  in  fact  to  account  for  the  toxic  effect  alone.  In  many 
instances  the  percentages  of  those  two  elements  found  were  as  high  in 
toxic  solutions  of  copper,  iron,  or  zinc  salts  as  when  toxic  concentra- 
tions of  calcium  or  magnesium  chlorides  were  used.  We  might, 
therefore,  in  the  light  of  our  present  knowledge,  be  justified  in  attrib- 
uting the  decreased  growth  of  the  plants  to  the  abnormally  high  ab- 
sorption of  calcium  and  magnesium  and  the  consequent  reactions 
taking  place  within  the  plant  cells.  The  permeability  of  the  mem- 
brane must  be  altered  to  allow  of  the  presence  of  these  ions  in  large 
numbers.  The  toxic  effects  due  to  the  presence  of  large  amounts  of 
calcium  or  magnesium  salts  might  be  evident  if  we  could  inject  solu- 
tions of  these  salts  into  the  plant  without  altering  the  permeability 
of  the  plasma  membrane.  But  from  the  present  data  it  seems  that 
the  alteration  in  the  permeability  of  the  membrane  is  the  essential 
consideration. 

It  is  probable  also  that  the  toxicity  of  any  solution  is  accompanied 
by  the  increased  permeability  of  the  plant  tissue  to  all  inorganic  salts 
which  are  normally  found  in  plants.  There  may  be  exceptions  as 
noted  already  for  iron  and  calcium,  but  in  general  this  relation  holds 
from  the  data  now  at  hand. 

Ruprecht61  has  localized  the  effects  of  aluminum  salts  in  the  few 
layers  of  cells  surrounding  the  root  hairs  and  attributes  the  death  of 
the  plants  grown  in  solutions  of  aluminum  salts  to  starvation  incident 
upon  the  inability  of  the  plant  to  obtain  nutrient  salts  for  normal 


ii  Mass.  Exp.  Sta.  Bull.  161   (1915),  p.  125. 


]918]  Way  nick:    Antagonism  and  Cell  Permeability  165 

metabolism.  Forbes62  has  likewise  localized  the  effects  of  copper  salts, 
when  present  in  toxic  concentrations,  and  concludes  that  the  toxic 
effect  of  copper  is  due  to  the  combination  of  metal  with  protein  at 
the  growing  tips  of  the  roots. 

From  the  experimental  results  given  in  the  present  paper,  it  is 
evident  that  the  presence  of  the  salts  of  each  element  in  toxic  concen- 
tration results  in  an  increased  permeability  of  the  plant  tissues  to 
calcium  and  magnesium  at  least.  Ruprecht's  view  that  plants  starve 
for  lack  of  nutrient  salts  when  grown  in  toxic  solutions  is  untenable, 
in  the  light  of  the  above  discussion. 

The  results  of  both  investigators  are  significant  in  indicating  the 
localization  of  the  effect  of  the  two  metals  studied  in  the  extreme 
outer  portion  of  the  roots,  in  which  the  plasma  membrane  is  located. 

The  results  obtained  by  Loeb  with  Fundulus  eggs,  by  Osterhout 
with  Laminaria,  using  electrical  conductivity  methods,  and  by  Brooks 
employing  microscopical  methods  with  various  plant  tissues,  all  point 
to  the  preservation  of  normal  permeability  as  the  result  of  antago- 
nistic salt  action.  The  results  reported  by  these  investigators  using 
widely  different  methods  have  been  confirmed  in  the  present  work  by 
the  use  of  a  more  direct  and  more  nearly  quantitative  method  than 
any  hitherto  employed. 

It  must  be  recognized,  however,  that  a  picture  of  but  one  stage  in 
the  growth  of  the  plant  has  been  given  and  that  only  a  portion  of  the 
inorganic  constituents  have  been  determined.  The  results  reported 
are  essentially  those  of  a  static  system  and  must  be  so  considered  in 
comparing  them  with  results  obtained  by  the  use  of  other  methods 
referred  to  above. 

Summary 

In  the  present  paper  results  are  given  showing  the  effect  of  vari- 
ous salt  solutions  upon  the  chemical  composition  of  plants,  with  spe- 
cial reference  to  a  correlation  between  toxic  and  antagonistic  effects 
and  composition.  A  uniform  nutrient  solution  was  used  throughout. 
The  cultures  were  arranged  in  series  in  which  the  concentration  of 
one  salt  was  kept  constant  while  the  concentration  of  a  second  salt 
varied  over  a  wide  range.  In  several  series  the  concentration  of  both 
varied,  but  the  ratio  between  the  two  remained  constant.  The  ana- 
lytical data  cover  the  percentages  of  calcium  and  magnesium  found 
in  the  plants  grown  in  every  culture,  together  with  determinations 


62  Univ.  Cal.  Publ.  Agr.  Sci.,  vol.  1  (1917),  p.  395. 


166  University  of  California  Publications  in  Agricultural  Sciences         [Vol.3 

of  potassium,  iron  and  copper  in  certain  series.  With  these  facts  in 
mind  the  results  of  the  investigation  may  be  briefly  stated  as  follows : 

The  composition  of  the  plants  grown  in  different  solutions  varied 
widely. 

Normal  growth,  i.e.,  approximately  that  of  the  controls,  was  always 
accompanied  by  approximately  equal  percentages  of  calcium  and 
magnesium  in  the  plants. 

In  nearly  all  cases  in  which  the  growth  of  the  plants  was  decreased 
to  a  marked  extent,  the  amounts  of  the  two  elements  referred  to  above 
were  increased  greatly. 

The  degree  of  absorption  of  any  salt  seems  to  be  independent  of 
the  concentration  present  in  the  solution  over  a  wide  range. 

Certain  relationships  are  pointed  out  between  calcium  and  mag- 
nesium absorption  and  the  presence  of  iron  and  zinc  salts  in  the 
solution. 

Antagonism  as  evidenced  by  growth  is  correlated  with  absorption 
of  the  ions,  which  were  determined,  in  every  instance. 

Stimulation  of  growth  was  recorded  when  ferric  sulphate  was 
present  in  the  nutrient  solution  in  certain  concentrations  and  with 
ferric  sulphate  and  zinc  sulphate  together. 

The  amounts  of  the  two  ions  uniformly  determined  were  not  neces- 
sarily found  in  the  same  proportions  in  roots  and  tops. 

The  possible  effects  of  changes  in  concentrations  of  the  various 
solutions  are  considered,  and  the  conclusion  reached  that  the  changes 
in  concentration  were  of  secondary  importance  over  the  range  of  con- 
centrations of  the  various  salts  used. 

Data  are  presented  showing  that  growth  is  the  same  with  widely 
varying  ratios  of  calcium  to  magnesium  in  the  nutrient  solution. 

The  results  in  general  confirm  those  of  Loeb,  Osterhout,  and 
Brooks  in  finding  that  antagonistic  salt  action  tends  toward  the 
preservation  of  normal  permeability  of  the  plasma  membrane  in  living 
tissue. 

This  problem  was  suggested  by  Dr.  C.  B.  Lipman.  The  writer 
wishes  to  express  his  thanks  for  this  and  for  many  other  valuable 
suggestions  offered  while  the  work  was  in  progress.  The  writer  is  also 
indebted  to  Prof.  L.  T.  Sharp  for  helpful  advice. 


1918]  Waynick:   Antagonism  and  Cell  Permeability  167 


NOTE 

The  following  key  applies  to  all  the  graphs.  The  numbers  on  the  abscissas 
represent  both  the  actual  weight  of  tops  and  roots  and  percentages  of  calcium 
and  magnesium,  or  of  iron,  when  the  latter  were  plotted.  The  numbers  on  the 
ordinates  correspond  to  the  number  of  cultures  as  given  in  the  table  on  the 
opposite  page.  The  heavy  lines  always  refer  to  the  roots,  the  light  lines  to 
the  tops. 

The  following  type  lines  are  used: 

(Solid  line)   Weight  of  tops. 

(Short  dashes)   Weight  of  roots. 

(Long  dashes)   Percentage  of  calcium. 

(One  long  and  two  short  dashes)   Percentage  of  magnesium. 

(One  long  and  one  short  dash)   Percentage  of  iron. 

The  numbers  given  in  the  "Explanation  of  Plates"  always  refer  to  the 
plants  arranged  in  order  from  left  to  right,  the  control  being  on  the  extreme 
right  in  every  case. 


168 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 


Table  1 
Calcium  Chloride 


No. 
1 


Solution 
CaCl2 

.002 


.004 


10 


11 


12 


Ful 


.01 


.02 


.04 


.06 


.08 


.10 


.12 


.16 


.20 


.24 


Dry  weight 

Tops   .7633 

.7104 

Roots  .3608 

Tops   .6486 

.4976 

Roots  .6555 

Tops   .6419 

.6138 

Roots  .5628 

Tops   .4814 

.6600 

Roots  .5750 

Tops   .5950 

.5442 

Roots  .5959 

Tops   .5692 

.4998 

Roots  .5048 

Tops   .6706 

.5015 

Roots  .5250 

Tops   .6114 

.4182 

Roots  .3827 

Tops   .4832 

.4778 

Roots  .3918 

Tops   .5668 

.5218 

Roots  .6123 

Tops   .5687 

.7637 

Roots  .6305 

Tops   .5266 

.5793 

Roots  .6067 


Nutrient  Tops   .7937 

.7418 

Roots  .6900 


.7368 
.1804 


.5731 
.3277 


.6279 
.2814 


.5707 

.2875 


.5696 
.2979 


.5845 
.2524 


.5860 
.2625 


.5148 
.1913 


.4805 
.1959 


.5443 
.3066 


.6662 
.3154 


.5529 
.3033 


.7677 
.3450 


I  « 

ft*  o 

15.38 
15.34 
26.24 

15.34 
17.34 
29.98 

17.29 
16.81 
28.10 

16.80 
17.69 
31.56 

17.52 
16.59 
33.54 

17.67 
17.00 
29.45 

13.92 
11.82 
29.46 

18.46 
19.94 
30.18 

16.80 
17.15 
29.12 

1 7.45 
18.47 
29.43 

17.03 
17.13 

31.82 

17.62 
17.34 

27.08 

18.80 
19.10 
20.03 


Pi 

15.36 


16.34 


17.05 


17.24 


17.05 


17.33 


12.87 


19.20 


16.97 


17.96 


17.08 


17.48 


18.95 


p4 

.477 


.231 

.400 
.470 
.261 

.514 
.517 
.219 

.504 
.450 
.121 

.640 
.565 
.113 

.718 
.713 
.098 

.407 
.684 
.363 

.428 
.440 
.290 

.443 
.425 
.273 

.387 
.486 
.392 

.348 
.388 
.204 

.354 
.392 
.373 

.393 
.390 
.227 


pi 

cS 
<D 

.477 


.435 


.515 


.477 


.602 


.715 


.545 


.434 


.434 


.436 


.368 


.373 


.391 


C  M 
g§ 
^=_ 

ft*  ° 

.051 
.046 
.413 

.116 
.154 
.507 

.095 

.078 
.498 

.045 
.044 
.483 

.048 
.044 
.563 

.050 

.087 
.420 

.035 
.079 
.472 

.061 
.068 
.652 

1.000 

.82 
.810 

1.340 

1.300 

.415 

.358 

.282 
.407 

.316 
.137 
.577 

.235 

.202 
.259 


a 

CS 
ID 

.049 


.135 


.086 


.045 


.046 


.069 


.057 


.064 


.962 


1.32 


.320 


.226 


.218 


Grown  October  24-December  5,  1915. 


1918] 


Wayniclc:   Antagonism  and  Cell  Permeability 


169 


Fig.  1 

Calcium  Chloride 

(See  Table  1) 


170  University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table 

2 

Magnesium  Chloride  4-  Calcium  Chloride 

Solution 

A 

Dry  Weight 

S3 

3 

0> 
g« 

3 

be 

a 

1  * 

go 

3 

a) 

c  fee 

Ph 

a 

0) 

<x> 

S  ® 

?    ° 

S3 

No. 

MgClo 

CaCl2 

4) 

3 

1 

.24 

.004 

Tops    .3407 

12.90 

.483 

1.21 

.3666 

.3536 

13.00 

12.95 

.500 

.491 

1.06 

1.16 

Eoots  .2456 

.1218 

20.05 

.557 

.670 

.022 

2 

.24 

.01 

Tops    .5106 

16.84 

.198 

.883 

.5862 

.5484 

16.88 

16.86 

.196 

.197 

.953 

.918 

Eoots  .3038 

.1519 

16.75 

.437 

.216 

.025 

3 

.24 

.02 

Tops    .2758 

15.22 

.490 

1.45 

.5885 

.4321 

15.94 

15.58 

.189 

.339 

1.09 

1.13 

Boots  .2533 

.1266 

21.02 

.407 

.433 

.010 

4 

.24 

.04 

Tops    .4450 

16.30 

.361 

.967 

.943 

.4828 

.4619 

.17.55 

16.92 

.342 

.351 

.920 

.943 

Eoots  .3018 

.1509 

16.32 

.550 

1.82 

.009 

5 

.24 

.06 

Tops    .3150 

18.50 

.445 

1.07 

.3716 

.3433 

17.87 

18.18 

.403 

.422 

1.13 

1.10 

Eoots  .2994 

.1497 

19.54 

.309 

.388 

.006 

6 

.24 

.08 

Tops    .6871 

15.97 

.154 

.440 

.6679 

.6775 

15.97 

.172 

.163 

.441 

.440 

Eoots  .4238 

.2119 

14.25 

.206 

.280 

.007 

7 

.24 

.10 

Tops    .4797 

16.75 

.458 

.645 

.4476 

.4636 

13.23 

14.99 

.457 

.457 

.507 

.576 

Eoots  .3000 

.1500 

19.95 

.430 

.577 

.002 

8 

.24 

.12 

Tops    .4583 

16.20 

.817 

.665 

.4279 

.4431 

17.43 

16.81 

.732 

.774 

.635 

.650 

Eoots  .2843 

.1421 

23.09 

.376 

.454 

.006 

9 

.24 

.16 

Tops    .3243 

15.14 

1.160 

1.46 

.3543 

.3393 

15.45 

15.29 

1.210 

1.18 

1.59 

1.52 

Eoots  .2276 

.1138 

24.60 

.970 

.714 

.010 

10 

.24 

.20 

Tops    .3404 

16.62 

1.140 

1.69 

.3132 

.3268 

17.50 

17.06 

1.330 

1.23 

1.98 

1.83 

Eoots  .2508 

.1254 

24.60 

1.200 

.785 

.012 

11 

.24 

.24 

Tops    .2707 

18.53 

1.640 

.807 

.015 

.2924 

.2815 

17.83 

18.18 

1.290 

1.46 

.826 

.816 

.009 

.012 

Eoots  .2107 

.1053 

24.62 

1.200 

.995 

.010 

12 

.24 

.30 

Tops    .4834 

18.48 

.563 

.675 

.010 

.5226 

.5030 

17.64 

18.06 

.585 

.574 

.707 

.691 

.012 

.012 

Eoots  .3888 

.1944 

25.16 

.437 

.538 

.010 

13 

.24 

Tops    .4281 

16.83 

.387 

.907 

.018 

.5106 

.4693 

17.20 

17.02 

.383 

.385 

.890 

.898 

.013 

.015 

Eoots  .2463 

.1231 

23.07 

.208 

.800 

.011 

Grown   January  2-February  13,  1916. 


1918] 


Waynick:    Antagonism  and  Cell  Permeability 


171 


0  .1   _ 


li 


13 


Fig.  2 

Magnesium  Chloride  +  Calcium  Chloride 

(See  Table  2) 


172 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table  3 
Magnesium  Sulphate  +  Calcium  Chloride 


Solution 


No. 
1 


9 

10 


1  1 


ll1 


MgCls 
.30 

.30 

.30 

.30 

.30 

.30 

.30 

.30 

.30 
.30 

.30 

.30 


13 

.30 

14 

.30 

Full  Nutrient 

CaCl2 
.001 

.002 

.004 

.01 

.02 

.04 

.06 

.08 

.10 
.12 

.16 

.20 
.24 
.30 


Dry  Weight 
Tops    .1986 
Eoots  .2650 

Tops    .2700 
Roots  .2766 

Tops    .2566 
Roots  .2236 

Tops    .3194 
Roots  .1744 

Tops    .2074 
Roots  .2249 

Tops    .5249 

.4166 

Roots  .1309 

Tops     .3036 

.1836 

Roots  .0850 

Tops    .2184 

.2403 

Roots  .0648 

Tops    .3978 
Roots 

Tops    .3842 

.2759 

Roots  .1059 

Tops    .8819 

.9600 

Roots  .6900 

Tops    .2100 
Roots  .2700 

Tops    .2576 
Roots  .1930 

Tops    .2600 
Roots  .2000 

Tops    .7937 

.7418 

Roots  .6900 


a 

cS 

.2318 

.2733 

.2401 

.2469 

.2163 

.4707 
.0654 

.2436 
.0425 

.2293 
.0324 


.3300 
.0529 

.9209 
.3450 

24.00 

.2253 

.2300 

.7677 
.3450 


16.60 
15.60 

16.44 
16.39 

18.43 
17.39 

16.49 
14.28 

14.56 
15.73 

15.19 
15.57 
18.10 

16.83 
17.42 
14.70 

17.90 
15.99 
13.05 

30.08 

17.92 
19.57 
27.29 

17.41 
18.11 
33.71 


18.80 
19.10 
20.03 


16.10 
16.41 
17.91 
15.38 
15.14 
15.38 

17.12 

16.94 

30.08 
18.74 

17.76 

14.08 
15.40 
14.30 

18.95 


.300 
.209 

.198 
.143 

.216 
.274 

.229 
.189 

.526 

.423 
.396 
.443 

.254 
.145 
.263 

.235 
.213 
.104 

.525 

.329 
.388 
.623 

.136 
.184 
.435 


.393 
.390 

2'27 


.259 
.170 
.255 
.209 
.526 
.409 

.254 

.224 

.525 
.358 

.160 

1.390 

1.490 

2.200 

.391 


M 

C3 

££ 

.440 
.413 

.571 
.514 

.595 
.533 

.515 

.088 

.749 
.565 

.405 
.437 


.649 

.790 

.700 
.663 
.230 

1.15 

.399 
.563 

.124 

.047 
.037 
.047 


.235 
.202 
.259 


.426 
.542 
.564 
.515 
.607 
.418' 

.719 

.681 

1.15 
.481 

.042 

3.660 
4.210 
3.790 

.218 


Grown  October  24  to  December  5,  1915. 


1918] 


WaynicTc:   Antagonism 


and  Cell  Permeability 


173 


•        chloride  +  Calcium  Chloride 
Magnesium  Chloride  -r 
8  (See  Table  3) 


174 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table  4 
Magnesium  Chloride  +  Calcium   Chloride 


Solution 

A 

Dry  Weight 

pi 

<v 

el 

03 

P-I 

CD 

No. 

MgCl2 

CaCl2 

CD    0 

1 

.36 

.02 

Tops 

.2878 

18.24 

.271 

1.71 

.3189 

.3033 

17.12 

17.68 

.311 

.291 

1.51 

Eoots 

.1644 

.0822 

18.15 

.221 

.98 

2 

.36 

.04 

Tops 

.4128 

18.25 

.374 

1.89 

.5210 

.4716 

16.55 

17.40 

.342 

.358 

1.99 

Eoots 

.3144 

.1572 

20.65 

.263 

1.24 

3 

.36 

.06 

Tops 

.2952 

16.31 

.431 

2.14 

.2640 

.2799 

17.21 

16.76 

.392 

.461 

2.37 

Roots 

.1560 

.0780 

17.90 

.472 

3.00 

4 

.36 

.08 

Tops 

.1878 

18.10 

.394 

2.22 

.2034 

.1999 

.2400 

21.05 

.407 

.400 

2.48 

Eoots 

.0684 

.0342 

17.80 

.733 

4.43 

5 

.36 

.10 

Tops 

.2292 

15.13 

.532 

2.90 

.1698 

.1999 

16.20 

15.66 

.614 

.573 

2.61 

Eoots 

.1272 

.0636 

17.90 

.631 

4.71 

6 

.36 

.12 

Tops 

.4788 

14.60 

.817 

.890 

.3938 

.4363 

13.65 

19.12 

.625 

.721 

1.02 

Eoots 

.2244 

.1122 

17.60 

.813 

3.33 

7 

.36 

.16 

Tops 

.3616 

14.12 

.832 

1.23 

.4410 

.4013 

15.17 

14.64 

.742 

.787 

1.72 

Eoots 

.2287 

.1143 

16.00 

.873 

2.47 

8 

.36 

.20 

Tops 

.2963 

13.21 

.931 

2.01 

.2505 

.2734 

12.17 

12.69 

.871 

.931 

2.32 

Eoots 

.1355 

.0677 

16.99 

1.020 

4.82 

9 

.36 

.24 

Tops 

.3100 

14.21 

.870 

2.31 

.2106 

.2603 

13.17 

13.69 

1.200 

1.03 

3.02 

Eoots 

.1734 

.0867 

14.20 

.713 

4.89 

Full  Nutrient 

Tops 

.7819 

20.40 

.349 

.223 

.7459 

.7639 

19.70 

20.05 

.330 

.339 

.268 

Eoots 

.6209 

21.70 

.242 

.248 

1.61 


1.94 


2.25 


2.35 


2.75 


.95 


1.47 


2.16 


.039 
.047 


.043 


.042 
.027 


.034 


.295 


Grown  January  7-February  21,  1916. 


1918] 


WaynicJc:   Antagonism  and  Cell  Permeability 


175 


1  .4 


1  . 


1.2  - 


1  .1 


1.0- 


0  .8 


0  .7 


0.6  - 


0  .5 


0.4  - 


0  .3 


0  .  2 


,'\ 


0.1  -  , 


Fig.  4 

Magnesium  Chloride  +  Calcium  Chloride 

(See  Table  4) 


176 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table 

;  5 

Magnesium 

Sulphate 

No. 

Solution 
MgS04 

Dry  Weight 

a 

a 
<v 

8 

M 

1 

.06 

Tops 

.7326 

16.78 

.7838 

.7587 

16.46 

16.62 

Roots 

.6383 

.3191 

16.84 

2 

.10 

Tops 

.7150 

15.73 

.9420 

.8276 

16.21 

15.97 

Roots 

.4822 

.2411 

22.91 

3 

.14 

Tops 

.6888 

15.09 

.6546 

.6717 

15.64 

15.36 

Roots 

.4437 

.2218 

20.31 

4 

.16 

Tops 

.3042 

12.65 

.3657 

.3349 

11.81 

12.23 

Roots 

.0762 

.0381 

21.52 

5 

.18 

Tops 

.2875 

12.32 

.2978 

.2926 

12.72 

12.52 

Roots 

.0542 

.0271 

20.49 

Full  Nutrient 

Tops 

.7819 

20.40 

.7459 

.7639 

19.70 

20.05 

Roots 

.6209 

21.70 

.358 
.379 
.495 

.302 
.313 

.580 

.201 
.230 

.148 

.337 
.331 
.985 

.496 

.371 

1.130 

.349 
.330 
.242 


.368 
.307 
.215 
.334 
.434 
.339 


.785 
.750 
.550 

.320 

.320 

1.270 

.350 
.390 
.740 

.350 
.310 
1.01 

.280 

.310 

1.300 

.223 

.268 

.248 


S3 

.767 


.320 


.370 


.330 


.295 


.295 


Grown  December  9-January  20,  1915-16 


1918] 


WaynicTc:   Antagonism  and  Cell  Permeability 


177 


1.4 


1.3- 


1.2  - 


1.1  - 


1.0  - 


0  .9 


0.8- 


0.7- 


0  .6 


0  .5 


0  .4 


0  .3 


0.2  - 


0.1- 


Fig.  5 

Magnesium  Sulphate 

(See  Table  5) 


178 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table 

;  6 

Magnesium  Chloride  +  Calcium   Chloride 

Solution 

A 

Dry  Weight 

3 

c 

too 
a 

C  eg 

go 

a 

CD 

CD    0 
P-l 

a 

0/ 

c 

C3 

No. 

CaCl2 

MgS04 

3 

1 

.04 

.18 

Tops 

1.1626 

19.91 

.261 

.407 

.036 

1.2526 

1.2076 

19.75 

19.83 

.332 

.296 

.722 

.564 

.023 

.030 

Eoots 

.9895 

.4942 

16.52 

.030 

.312 

.017 

2 

.08 

.18 

Tops 

1.1888 

20.43 

.312 

.077 

.060 

1.0948 

1.1418 

21.15 

20.79 

.377 

.344 

.084 

.081 

.060 

.060 

Eoots 

.9037 

.4518 

26.13 

.151 

.180 

.020 

3 

.12 

.18 

Tops 

1.1287 

19.50 

.275 

.021 

.024 

1.2691 

1.1989 

21.22 

20.36 

.276 

.275 

.023 

.022 

.021 

.023 

Eoots 

.9591 

.4795 

20.93 

.560 

.185 

.020 

4 

.16 

.18 

Tops 

1.2293 

17.85 

.169 

.035 

.036 

1.2134 

1.2213 

16.75 

17.30 

.131 

.150 

.049 

.042 

.036 

Eoots 

.5444 

.2722 

16.47 

.690 

.101 

.024 

5 

.20 

.18 

Tops 

1.1786 

21.67 

.333 

.039 

.033 

1.1021 

1.1403 

21.07 

21.37 

.256 

.294 

.044 

.041 

.033 

Eoots 

1.0500 

.5250 

25.40 

.820 

.299 

.028 

6 

.24 

.18 

Tops 

.9773 

21.47 

.448 

.462 

.045 

.9923 

.9848 

21.65 

21.56 

.545 

.496 

.486 

.470 

.033 

.039 

Eoots 

.8600 

.4300 

26.07 

1.290 

.407 

.024 

7 

.28 

.18 

Tops 

.8459 

18.62 

.476 

.437 

.032 

.7450 

.7954 

20.05 

19.33 

.523 

.499 

.409 

.420 

.037 

.035 

Eoots 

.6014 

.3007 

24.87 

1.580 

.565 

.023 

8 

.32 

.18 

Tops 

.4608 

21.43 

.640 

1.05 

.027 

.5316 

.4962 

21.43 

.568 

.604 

1.24 

1.14 

.019 

.023 

Eoots 

.3350 

.1675 

29.00 

3.330 

1.07 

.026 

9 

.36 

.18 

Tops 

.1916 

18.56 

.747 

1.43 

.048 

.2349 

.2132 

18.80 

18.68 

.955 

.851 

1.59 

1.50 

.049 

.047 

Eoots 

.0976 

.0488 

18.05 

1.000 

2.80 

.2269 

.1819 

13.60 

13.50 

.235 

.370 

1.71 

1.71 

.031 

.053 

10 

.18 

Tops 
Eoots 

.1369 
.0614 

.0307 

13.40 
16.62 

.505 
.112 

2.03 
3.68 

.016 

Grown  January  24-March  6,  1916. 


1918] 


Waynick:   Antagonism  and  Cell  Permeability 


179 


1  .  4 


1  .3 


1  .  2 


1  .  1 


1  .0 


0  .9 


0.8   - 


0  .7 


0.6   - 


0  .5 


0.4" 


0  .3 


0.2    " 


0  .  1 


Fig.  6 

Magnesium  Sulphate  +  Calcium  Chloride 

(See  Table  6) 


180 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table  7 
Potassium  Chloride 


Solution 


fc    Ca(N03)2  MgSO*       Dry  Weight 


3 


a 


.04 


.08 


.12 


.16 


.20 


.24 


.28 


.15 


.15 


.15 


.15. 


.15 


Tops    1.8850 

1.1941 

Eoots     .9776 

Tops    1.7825 

1.5648 

Roots     .8813 


17.90 
1.5395     20.00     18.95 
.4588     19.35 

16.20 
1.6731     16.92     16.56 
.4406     25.37 


.15       Tops    1.6850 

Eoots     .6284 
.15       Tops    1.2078 

Roots     .5668 


Tops 
Roots 


19.90 

.8425      

.3141     31.20 

16.67 

.6030      

.2834     27.80 

No  growth 


19.90 


.067 
.067 
.062 

.101 
.120 
.135 

.487 

.037 
.247 

.037 


.967 

.067       .808 

.101 

.762 

.110       .842 

.430 


•Z        g 


.487 


.24' 


2.190 

2.390 
2.120 


2.410 


.802 


2.190 


2.120 


Tops      .5818 
Roots     .1581 


Tops 
Roots 


8    Full  Nutrient    Tops    1.5682 

1.5775 

Roots     .9776 


.026 
.018 
.093 


.022 


.024 

.021     .023 

.012 


18.50 

.550 

9.20 

o 
o 

.2909      

18.50 

.550 

9.200 

-«j 

.0740     21.90 

.195 

6.520 

§ 

o 

s 

No  growth 

<! 

18.33 

.293 

.231 

1.5728     19.40 

18.86 

.312 

.302 

.279 

.255 

20.40 

.271 

.279 

Grown  April  14-May  27,  1916. 


1918] 


Waynick:   Antagonism  and  Cell  Permeability 


181 


0.9  - 


Fig.  7 

Magnesium  Sulphate  -\-  Calcium  Nitrate 

(See  Table  7) 


182 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 


Table  8 
Magnesium   Sulphate  +  Calcium   Nitrate 


No. 

Solution 
KC1         Dry  Weight 

3 

0)   o 

P4 

3 

pi 

03 

3 

fe«H 

Ph 

3 

fe«H 

Ph 

pi 

CD 
3 

1 

.04 

Tops 

1.1563 

19.65 

.078 

.228 

.028 

1.37 

1.1128 

1.1345 

19.65 

19.65 

.201 

.214 

.045 

.037 

1.80 

1.58 

Eoots 

.5038 

.2519 

17.07 

.256 

.175 

.173 

.61 

2 

.06 

Tops 

1.0773 

24.00 

.128 

.083 

.041 

3.54 

1.0791 

1.0782 

25.90 

24.95 

.113 

.120 

.118 

.100 

.030 

.036 

3.54 

3.54 

Boots 

.5979 

.2989 

16.27 

.241 

.170 

3 

.08 

Tops 

.7638 

27.50 

.302 

.201 

.064 

4.81 

4.56 

Eoots 

.4659 

.1829 

17.20 

.238 

.180 

.106 

4.80 

4 

.10 

Tops 

.8417 

25.90 

.204 

.302 

.268 

3.56 

.7900 

.8158 

24.40 

25.15 

.186 

.195 

.205 

.253 

.280 

.274 

3.80 

3.68 

Eoots 

.3135 

.1567 

14.20 

.570 

.300 

.072 

1.02 

5 

.12 

Tops 

.5656 

31.00 

.283 

.378 

.195 

4.78 

.5815 

.5734 

29.80 

30.40 

.265 

.274 

.657 

.567 

.072 

.133 

4.30 

4.54 

Eoots 

.2215 

.1107 

18.00 

.780 

.101 

.175 

1.67 

6 

.14 

Tops 

.4825 

39.50 

.217 

.478 

.091 

9.11 

.4200 

.4531 

32.30 

35.90 

.222 

.219 

.454 

.466 

.092 

.091 

7.10 

8.10 

Eoots 

.1762 

20.83 

.617 

.139 

.021 

1.53 

7 

.16 

Tops 

.5375 

.2687 

42.00 

42.00 

.248 

.248 

.730 

.730 

.071 

.071 

11.21 

11.21 

Eoots 

.1339 

.0669 

23.83 

.505 

1.34 

.250 

2.31 

Grown  January  31-March  13,  1916. 


1918] 


WaynicTc:   Antagonism  and  Cell  Permeability 


183 


1  .  2 


1  .  1 


1  .  0 


0.9- 


0.8- 


0  .7 


0.6- 


0.5  - 


0.4   - 


0.3  - 


0.2^ 


0  .1 


7>/M 


■/ 


v--r 


4 


Fig.  8 

Potassium  Chloride 

(See  Table  8) 


184  University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 


Table  9 
Magnesium  Sulphate  +  Potassium  Chloride 


Solution 

A 

Drv  Weight 

a 

03 

3 

bfl 

s -a 

a 

So 

c3 

i  * 

CD    0 

P-l 

o3 

OJ 

3 

03 

C3 

p-i 

03 

C 

03 

No. 

MgS04 

KC1 

Oj 

3 

1 

.04 

.18 

Tops 

.6579 

19.65 

.141 

.490 

2.13 

.6379 

.6479 

21.20 

20.42 

.162 

.151 

.500 

.495 

1.72 

1.92 

Eoots 

.2800 

.1400 

20.00 

2.79 

.67 

2 

.08 

.18 

Tops 

.6800 

22.07 

.138 

.430 

3.21 

.7162 

.6981 

25.10 

23.58 

.171 

.154 

.480 

.455 

3.00 

3.10 

Boots 

.2682 

.1341 

23.50 

.590 

1.350 

.72 

3 

.12 

.18 

Tops 

.4957 

22.60 

.292 

3.78 

.5202 

.5079 

26.50 

24.55 

.310 

.301 

.923 

.923 

3.82 

3.80 

Roots 

.2200 

.1100 

25.25 

.745 

.104 

.86 

4 

.16 

.18 

Tops 

.6229 

23.50 

.152 

.382 

4.12 

.6464 

.6346 

24.70 

24.10 

.169 

.161 

.381 

.381 

d 

4.21 

4.16 

Eoots 

.3123 

.1561 

22.70 

.291 

.561 

f-{ 

1.40 

5 

.20 

.18 

Tops 

.3643 

.4819 

.4231 

26.20 

26.20 

.440 

.440 

.977 

.977 

03 
O 

5.01 

5.01 

Roots 

.2063 

.1031 

24.00 

.678 

1.060 

4J 

2.20 

6 

.24 

.18 

Tops 

.3259 

33.20 

.407 

1.040 

7.22 

.2141 

.2700 

37.60 

35.40 

.423 

.415 

2.920 

1.970 

O 

o 

8.13 

7.66 

Roots 

.1297 

.0648 

22.10 

.750 

1.490 

+3 

2.80 

7 

.28 

.18 

Tops 

.1522 

31.10 

.299 

1.030 

o 

9.20 

.2050 

.1786 

35.10 

33.10 

.265 

.282 

.990 

1.010 

a 

11.30 

10.25 

Roots 

.0913 

.0456 

24.10 

• 

.735 

2.900 

< 

3.10 

8 

.32 

.18 

Tops 

.2704 

36.10 

.263 

.890 

11.20 

.2997 

.2850 

36.20 

36.15 

.211 

.237 

.810 

.850 

7.20 

9.20 

Roots 

.1467 

.0733 

22.57 

.229 

.818 

2.17 

9 

.36 

.18 

Tops 

.2800 

23.90 

.261 

.801 

3.21 

.2846 

.2823 

25.00 

24.45 

.278 

.269 

.895 

.848 

4.17 

3.64 

Roots 

.1417 

.0708 

21.60 

.699 

.248 

2.18 

Fu] 

11  Nutrient 

Tops 

1.0992 

20.17 

.310 

.268 

1.0750 

1.0872 

19.12 

18.69 

.297 

.303 

.228 

.224 

Roots 

.8120 

20.00 

.271 

.233 

Grown  January  31-March  13,  1916. 


191S 


Waynick:   Antagonism  and  Cell  Permeability 


185 


1  .4 


1.3- 


1  .  2 


1  .  1 


1  .  0 


0  .  9 


0.8- 


0  .  7 


0  .  6 


0  .  5 


0.4- 


0.3- 


0  .  2 


0  .1 


Fig.  9 

Magnesium  Sulphate  +  Potassium  Chloride 

(See  Table  9) 


186 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 


Table  10 
Aluminum  Chloride 


£S 


No. 

Solution 
A1C13 

Dry  Weight 

P4 

c8 

3 

3 

Ph 

03 

3 

03 

1 

.0000033 

Tops 

.4438 

22.42 

.098 

.450 

.075 

.4688 

.4563 

20.40 

21.40 

.092 

.095 

.590 

.520 

.077 

.076 

Eoots 

.4315 

.2157 

20.20 

.055 

.590 

.089 

2 

.0000165 

Tops 

.2934 

23.20 

.196 

.793 

.151 

.2745 

.2839 

22.70 

22.95 

.180 

.188 

.793 

.793 

.140 

.145 

Roots 

.1976 

.0988 

18.70 

.250 

.990 

.061 

3 

.000033 

Tops 

.2872 

21.09 

.316 

.953 

.173 

.3595 

.3233 

22.60 

21.84 

.321 

.318 

1.000 

.970 

.154 

.163 

Eoots 

.2964 

.1482 

19.55 

.153 

.504 

.130 

4 

.000066 

Tops 

.3028 

21.09 

.342 

.605 

.128 

.3200 

.3114 

22.80 

21.94 

.334 

.338 

.615 

.610 

.155 

.141 

Roots 

.2422 

.1211 

19.55 

.334 

.840 

.171 

5 

.000132 

Tops 

.2720 

22.60 

.423 

.835 

.101 

.2479 

.2599 

19.00 

20.80 

.450 

.436 

.837 

.836 

.134 

.117 

Roots 

.2645 

.1322 

26.60 

.382 

1.110 

.104 

6 

.000331 

Tops 

.3178 

20.20 

.246 

1.130 

.137 

.4016 

.3597 

21.90 

21.05 

.247 

.246 

.690 

.91 

.124 

.130 

Roots 

.2550 

.1275 

27.90 

.108 

.108 

7 

.00331 

Tops 

.3863 

22.30 

.258 

.535 

.114 

.2369 

.3116 

21.82 

22.60 

.268 

.263 

1.230 

.88 

.187 

.150 

Roots 

.2153 

.1076 

29.40 

.583 

.330 

.231 

Grown  August  26-October  7,  1916. 


1918] 


Wayniclc:   Antagonism  and  Cell  Permeability 


187 


1  .2 


1  .  1 


1  .0 


0.9  - 


0  .7 


0.6- 


0  .5 


0  .  4 


0  .3 


0  .2 


0  .1 


I  I 


Fig.   10 

Aluminum  Chloride 

(See  Table  10) 


188 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table  11 


No. 

1 


Cal 

cium  C 

!hlorid( 

i  4-  Aluminum 

Chloride 

Solution 

A 

Dry  Weight 

03 

u 

es 

I" 
®  0 

3 

Percentage 
of  Ca 

Mean 

0 

03 
C   fcJD 

03 

3) 

3 

03 

M 
03 

a 

A1C13 

CaCl2 

.0000033 

.20 

Tops 

.2962 

24.20 

1.630 

.745 

.2087 

.2524 

23.10 

23.65 

1.820  1.72 

.830 

.787 

Eoots 

.1897 

.0948 

20.40 

.730 

.872 

.0000165 

.20 

Tops 

.2542 

24.23 

1.610 

.890 

.123 

.2678 

.2610 

22.18 

23.2'0 

1.720  1.66 

9.30 

.910 

.313 

.21 

Eoots 

.2745 

.1372 

26.17 

.731 

.0000331 

.20 

Tops 

.3324 

23.00 

1.250 

.787 

.503 

.3706 

.3515 

25.20 

24.10 

1.460  1.35 

.903 

.845 

.540 

.52 

Roots 

.2942 

.1471 

26.10 

.790 

.817 

.075 

.0000662 

.20 

Tops 

.2139 

23.20 

1.210 

.821 

.237 

.2337 

.2238 

21.70 

22.45 

1.030  1.12 

.733 

.772 

.114 

.17 

Eoots 

.1839 

.0919 

24.20 

.621 

1.02 

.000132 

.20 

Tops 

.2148 

20.58 

1.400 

.932 

.248 

.18 

.3085 

.2611 

24.10 

22.34 

1.060  1.23 

.897 

.913 

.119 

.18 

Eoots 

.2239 

.1119 

20.20 

.513 

.947 

.000331 

.20 

Tops 

.2337 

24.10 

1.420 

.632 

.105 

.2137 

.2237 

21.32 

22.71 

1.270  1.34 

.711 

.621 

.210 

.15 

Eoots 

.1682 

.0841 

20.16 

.672 

1.310 

00331 

.20 

Tops 

.2496 

24.10 

1.330 

.490 

.155 

.1946 

.2221 

22.90 

23.50 

1.550  1.44 

.516 

.503 

.285 

.22 

Eoots 

.1296 

.0648 

26.40 

.780 

.253 

.021 

Nutrient 

Tops 

.7103 

18.17 

.322 

.221 

.7327 

.7215 

19.31 

18.74 

.377     .349 

.270 

.245 

Eoots 

.6600 

.3300 

19.99 

.300 

.233 

Crown  August  26-October  7,  1916 


1918] 


IV ay  nick:    Antagonism  and  Cell  Permeability 


189 


1  .4 


1  .3 


1  .  2 


1.1  - 


1  .0 


0.9  - 


0.8  - 


0.7- 


0.6  - 


0.5  - 


0  .4 


0.3  - 


0.1  - 


Fig.  11 

Calcium  Chloride  +  Aluminum  Chloride 

(See  Table  11) 


190 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 


Table  12 
Aluminum  Chloride  +  Magnesium  Chloride 


Solution 

A 

Dry  Weight 

03 

Ph 

03 

a 

3 

fe«H 

Pi 

03 

3 

03 

No. 

A1C13 

MgCl2 

3 

1 

.0000033 

.20 

Tops 

.2819 

19.02 

.426 

2.13 

.121 

.5457 

.4138 

18.71 

18.86 

.371 

.398 

1.91 

2.02 

.131 

.126 

Eoots 

.2936 

.1468 

21.31 

.321 

2.12 

.122 

2 

.0000165 

.20 

Tops 

.3539 

17.31 

.327 

1.71 

.141 

.3000 

.3269 

19.21 

18.16 

.421 

.374 

1.83 

1.77 

.132 

.136 

Eoots 

.1750 

.0875 

22.17 

.400 

1.71 

.101 

3 

.0000331 

.20 

Tops 

.2243 

21.05 

.352 

2.51 

.148 

.2259 

.2251 

17.70 

19.39 

.448 

.400 

2.80 

2.65 

.147 

.147 

Eoots 

.1239 

.0614 

24.10 

.366 

1.96 

.135 

4 

.0000662 

.20 

Tops 

.6327 

19.00 

.147 

1.46 

.050 

.7644 

.6985 

18.10 

18.55 

.130 

.138 

1.36 

1.42 

.047 

.049 

Eoots 

.5526 

.2763 

19.95 

.111 

.333 

.040 

5 

.000132 

.20 

Tops 

.2406 

19.75 

.280 

2.78 

.069 

.3156 

.2781 

19.55 

19.65 

.282 

.281 

3.11 

2.94 

.064 

.066 

Eoots 

.1250 

.0625 

24.00 

.186 

5.88 

.181 

6 

.000331 

.20 

Tops 

.2279 

16.50 

.407 

.940 

.091 

.2048 

.2163 

19.90 

18.20 

.467 

.437 

.557 

.74 

.081 

.086 

Eoots 

.1700 

.0850 

22.76 

.302 

2.51 

.098 

7 

.00331 

.20 

Tops 

.2400 

17.40 

.131 

2.14 

.161 

.1450 

.1925 

18.80 

18.10 

.161 

.145 

1.71 

1.92 

.192 

.176 

Eoots 

.0990 

.0495 

32.60 

.200 

4.53 

.439 

Grown  August  26-October  7,  1916. 


1918] 


JVaynicJc:   Antagonism  and  Cell  Permeability 


191 


1.4- 


1.3- 


1  .2 


1.1- 


1.0  - 


0.9 


0.8 


0.7  - 


0  .6 


0.5  - 


0.4- 


0  .3 


0  .2 


0.1  - 


\^  - 


I  I  I  I  I 

12  3  4  5 


Fig.  12 

Aluminum  Chloride  +  Magnesium  Chloride 

(See  Table  12) 


192 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table  13 
Ferric  Chloride 


No. 

Solution 
FeCl3 

Dry  Weight 

3 

Ph 

3 

So 

Ph 

Ph 

c 

5!     O) 
£^ 

Sh«w 

Ph 

3 

1 

.000089 

Tops 

.7712 

14.50 

.199 

.258 

.214 

.8376 

.8044 

19.20 

16.85 

.228 

.213 

.237 

.247 

.199 

.206 

Roots 

.7143 

.3571 

2 

.000168 

Tops 

.7884 

17.90 

.138 

.244 

.200 

1.2762 

1.0323 

19.30 

16.60 

.106 

.122 

.299 

.271 

.197 

.198 

Roots 

.6462 

.3231 

17.60 

.178 

.250 

.220 

3 

.00168 

Tops 

1.3519 

15.70 

.045 

.259 

.247 

1.1269 

1.2394 

18.10 

16.90 

.046 

.045 

.225 

.242 

.232 

.239 

Roots 

.9034 

.4517 

17.10 

.054 

.213 

4 

.0168 

Tops 

1.0000 

16.78 

.033 

.254 

.109 

1.2750 

1.1370 

16.51 

16.64 

.028 

.030 

.222 

.238 

.138 

.123 

Roots 

.7219 

.3609 

20.98 

.068 

.203 

.231 

Full  Nutrient 

Tops 

1.0992 

20.17 

.310 

.268 

1.0750 

1.0872 

19.12 

18.69 

.297 

.303 

.228 

.224 

Roots 

.8120 

20.00 

.271 

.233 

Grown  January  24-March  6,  1916. 


1918] 


Waynick:   Antagonism  and  Cell  Permeability 


]93 


1  .  4 


1  .3 


1.1- 


1  .0 


0  .9 


0  .7 


0.6 


0  .5 


0  .4 


0  .3 


0  .2 


0  .1 


v^—-"~- 


i  i  i  i 


Fig.  13 
Ferric  Chloride 
(See  Table  13) 


194 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 


Table  14 
Ferric  chloride  4-  Calcium  Chloride 


Solution 

A, 

Dry  Weight 

'a 
3 

0-3 
Ph 

3 

go 

No. 

FeCl3 

CaClo 

PM 

1 

.000089 

.20 

Tops 

.4826 

18.27 

1.02 

.4116 

.4471 

18.50 

18.38 

1.83 

Eoots 

.4143 

.2071 

28.35 

.531 

2 

.000168 

.20 

Tops 

.5238 

17.72 

1.77 

.5847 

.5592 

16.04 

16.88 

1.01 

Eoots 

.6115 

.3057 

22.70 

.188 

3 

.000352 

.20 

Tops 

.2732 

20.42 

2.78 

.3204 

.2918 

20.61 

20.51 

3.28 

Eoots 

.1774 

.0887 

22.40 

1.76 

4 

.000712 

.20 

Tops 

.5910 
.3810 

.4860 

19.01 

19.01 

2.85 
2.15 

Eoots 

.5053 

.2526 

31.30 

1.06 

5 

.00142 

.20 

Tops 

.5427 

16.56 

1.06 

.6353 

.5890 

17.79 

17.17 

1.33 

Eoots 

.6238 

.3119 

29.50 

.350 

6 

.00356 

.20 

Tops 

.2715 

14.76 

1.32 

.2632 

.2673 

17.09 

15.92 

1.23 

Eoots 

.2196 

.1098 

20.03 

1.13 

7 

.0058 

.20 

Tops 

.1792 

20.03 

2.37 

.2514 

.2153 

20.92 

20.47 

2.08 

Eoots 

.0636 

.0318 

29.30 

1.68 

8 

.0168 

.20 

Tops 

.3293 

17.03 

.584 

.4393 

.3843 

19.08 

18.05 

.460 

Eoots 

.3565 

.1783 

28.04 

.505 

«4H  ^ 


1.42 


1.39 


3.03 


2.00 


1.19 


1.27 


2.22 


.522 

Grown  December  9-January  19,  1916. 
Tron  determined  colorimetrically  in  this  series. 


.164 
.137 
.222 

.194 
.101 
.229 

.256 
.255 
.692 

.212 
.262 
.026 

.064 
.051 
.010 

.097 
.133 
.018 

.256 
.297 
.020 

.246 
.175 
.049 


.150 


.147 


.255 


.237 


.058 


.115 


.276 


.210 


.100 

.080 
.090 
.03 


.085 


.370 

.400     .38 
.120 


.50 
.40 
.02 

.36 
.50 
.16 


.45 


.43 


1.18 

1.02     1.10 
.98 


.800 
.900 
.200 

.148 
.120 
.06 


.134 


191S] 


WaynicJc:   Antagonism  and  Cell  Permeability 


195 


1.4- 


1  .3 


1.2- 


1  .1 


1  .0 


0  .9 


0  .  8 


0  .7 


0.6- 


\ 


/  \ 


0  .  5 


0  .4 


Fig.   14 

Ferric  Chloride  +  Calcium  Chloride 

(See  Table  14) 


106 


University  of  California  Publications  in  Agricultural  Sciences 


I  Vol.  3 


Table  15 
Ferric   Chloride  +  Magnesium  Chloride 


Solutic 

A 

in 

Dry  Weight 

03 

13 

a 

jo 

n 

03 

C  be 
03   O 

C    03 

Ah 

0 
c3 

No. 

FeCls 

MgCls 

3 

1 

.0000S9 

.20 

Tops 

.9650 

18.48* 

.182 

.874 

.091 

.7444 

.8547 

18.70 

19.59 

.163 

.172 

.833 

.853 

.111 

.101 

Eoots 

.6126 

.3063 

25.05 

.204 

.547 

.047 

2 

.000168 

.20 

Tops 

.7875 

18.12 

.181 

.189 

.210 

.9525 

.8700 

18.67 

18.44 

.164 

.172 

.191 

.190 

.210 

Eoots 

.6816 

.3408 

21.40 

.229 

.553 

.033 

3 

.000352 

.20 

Tops 

.8787 

15.17 

.172 

.218 

.067 

.7365 

.8076 

17.15 

16.16 

.137 

.154 

.310 

.264 

.073 

.070 

Eoots 

.6525 

.3262 

22.95 

.176 

.515 

4 

.000712 

.20 

Tops 

1.0114 

16.77 

.324 

.116 

.093 

1.1515 

1.1464 

17.57 

17.17 

.311 

.317 

.119 

.117 

.094 

.093 

Eoots 

.4837 

.2418 

21.86 

.258 

.503 

.057 

5 

.00142 

.20 

Tops 

.6648 

17.40 

.209 

.069 

.233 

.7927 

.7287 

16.22 

16.81 

.185 

.197 

.103 

.086 

.209 

.221 

Eoots 

.4627 

.2313 

22.12 

.258 

.093 

.058 

0 

.00356 

.20 

Tops 

1.1639 

16.50 

.176 

.153 

.147 

.9476 

1.0557 

16.10 

16.30 

.109 

.142 

.153 

.128 

.137 

Eoots 

.3298 

.1649 

20.28 

.433 

.936 

.066 

7 

.0058 

.20 

Tops 

.6608 

17.08 

.340 

.218 

.040 

.6189 

.7398 

16.07 

16.57 

.400 

.370 

.234 

.226 

.040 

Eoots 

.4846 

.2423 

20.09 

.235 

.788 

.050 

8 

.0168 

.20 

Tops 

1.0927 

18.96 

.344 

1.020 

.140 

1.0824 

1.0875 

20.05 

19.45 

.391 

.367 

1.100 

1.06 

.120 

.130 

Eoots 

.6358 

.3179 

21.05 

.177 

.262 

.051 

Grown  January  24-March  6,  1916. 


1918] 


Waynick:    Antagonism  and  Cell  Permeability 


197 


1.2  - 


1  .1 


1  .0 


o  .  y 


o  .  8 


0.7- 


0  .6 


0.5   - 


0  .4 


0.3- 


0  .  2 


0  .  1 


\ 


7 


^  All 


y.\  \ 


& 


Fig.  15 

Ferric  Chloride  +  Magnesium  Chloride 

(See  Table  15) 


198 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table  16 

Copper 

Chlori 

de 

No. 

Solution 
CuCl2 

Drv  Weight 

3 

CD 
CS 

U  54_| 

eg 

3 

fcJO 

Ph 

CD 

3 

CO 

bfi 

#?  ° 

3 

03 
M 

S3 

Pn 

3 

CD 
h£ 

"S  pi 

Ph 

c6 
CO 

1 

.000038 

Tops 

.4611 

25.60 

.425 

.194 

.120 

Boots 

.4692 

.4651 

23.60 

24.60 

.420 

.412 

.191 

.192 

.201 

.160 

2 

.000079 

Tops 

.2492 

20.20 

.455 

.716 

.155 

Eoots 

.3242 

.2867 

24.12 

22.16 

.563 

.509 

.723 

.719 

.221 

.188 

3 

.00015 

Tops 

.3887 

24.50 

.312 

1.10 

.114 

Eoots 

.4198 

.4042 

25.30 

24.90 

.361 

.336 

.90 

1.00 

.0261 

.070 

4 

.00031 

Tops 
Roots 

.2744 

.1372 

18.45 

18.45 

.150 

.150 

1.43 

1.48 

.040 

.040 

.001 

5 

.00047 

Tops 

.2581 

22.00 

.176 

1.35 

.214 

.002 

Roots 

.2344 

.2462 

18.10 

20.05 

.102 

.139 

1.02 

1.18 

.094 

.154 

.002 

.002 

6 

.00063 

Tops 

.0785 

19.10 

.254 

3.77 

.003 

Roots 

.0700 

.0742 

18.21 

18.65 

.425 

.339 

5.10 

4.43 

.248 

.248 

.005 

.004 

7 

.00079 

Tops 
Roots 

No 

growth. 

8 

.00198 

Tops 

Roots 

No 

growth. 

9 

.00392 

Tops 
Roots 

No 

growth. 

Full 

Nutrient 

Tops  1.1234 

19.20 

.311 

.213 

1.0268 

1.0751 

21.00 

20.10 

.241 

.276 

.199 

.206 

Roots 

.7210 

18.99 

.299 

.216 

Grown  March  9-April  20,  1916. 


1918] 


Waynick:   Antagonism  and  Cell  Permeability 


199 


1.4- 


1  .3 


1  .2 


1  .  1 


1.0- 


0.9  - 


0  .8 


0.7- 


/        \ 


\  / 


/ 


0  .6 


0.5  - 


0.4  - 


0  .3 


0  .1 


Fig.  16 

Copper  Chloride 

(See  Table  16) 


200 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.  3 


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191S] 


Way  nick:    Antagonism  and  Cell  Permeability 


201 


1.4- 


1.3- 


1.2- 


1.1  - 


1.0  - 


0.9- 


0.8- 


0.7   - 


0  .  6 


0  .5 


0.4- 


0  .3 


0.2- 


0  .1 


Fig.  17 

Copper  Chloride  +  Ferric  Chloride 

(See  Table  17) 


202 


University  of  California  Publications  in  Agricultural  Sciences       [Vol.  3 


Table  18 
Mercuric  Chloride  +  Copper  Chloride 


Solution 


No.         HgClo             CuCl2            Dry  Weight  3           PM                E 

1  .0000047     .0000023     Tops    .1175  20.15 

.1608  .1392  21.20     20.67 

Eoots  .0869  .0434  18.65 

2  .0000094     .0000047     Tops    .1792  20.00 

.2675  .2233  20.32     20.16 

Roots  .0846  .0423  16.71 


24.30 


3 

.0000184 

.0000094 

Tops 

.3778 
.3328 

.3503 

24.70 
23.30 

Roots 

.0928 

.0467 

15.45 

4 

.000047 

.000023 

Tops 

.3600 
.2334 

.2967 

21.22 
23.20 

Roots 

.1109 

.0554 

17.32 

5 

.000094 

.000047 

Tops 

.3643 
.4372 

.4007 

22.00 
20.12 

Roots 

.1109 

.0554 

17.32 

6 

.000189 

.000094 

Tops 

.2385 
.3105 

.2745 

22.30 
24.71 

Roots 

.1146 

.0573 

12.71 

7 

.000378 

.000188 

Tops 

.1500 
.2150 

.1825 

21.60 
23.40 

Roots 

.0609 

.0304 

12.82 

22.21 


21.06 


23.50 


22.50 


§5  =* 

Ph 

3 

££ 

Ph 

13 

3 

C  a 
Pu 

.674 

1.21 

.517 

.595 

.935 

1.07 

1.220 

.276 

.531 

.§73 

.612 

.572 

.942 

.907 

1.410 

.291 

.493 

.488 

.3 

.570 

.531 

.488 

1.360 

.211 

0) 

^5 

.431 

.834 

O 

.521 

.471 

.921 

.877 

'A 

.359 

.361 
.737 

o 

.354 

.356 

.695 
.361 

.716 

13 

DO 

.521 

.921 

o 
o 
H 

.503 

.512 

.973 

.947 

1.340 

.333 

.573 

1.212 

.672 

.622 

1.071 
.412 

1.14 

Grown  January  24-March  6,  1916. 


1918] 


TV  ay  nick-:    Antagonism  and  Cell  Permeability 


203 


1.4- 


1  .3 


1  .  2 


1  .  1 


1.0  - 


0.9  - 


0.8 


0.6  - 


0.5  - 


0  .4 


0.3  - 


0  .2 


0.1  - 


/^ 


/ 


\ 


'  I 


I  I 


Fig.  18 

Mercuric  Chloride 

(See  Table  18) 


204 


University  of  California  Publications  in  Agricultural  Sciences       [Vol.  3 


Table  19 
Copper  Sulphate 


Solution 
Xo.         CuS04 

1   .0000048 


3   .0000188 


4   .0000378 


5  .0000567 


S  S2        * 

Dry  Weight  3  fc  3 

Tops       .6148  22.80 

.7726     .6937  21.71     22.25 

Eoots     .4406     .2203  20.61 


.0000094      Tops 


SO 

Ph 

.104 

.117 

.153 


5394  21.42  .066 

.8462  .6928  17.50  17.46       .056 

Eoots     .4512  .2256  16.40 

Tops       .4712  24.40  .177 

.3128  .3920  18.90  21.65       .234 

Koots     .2948  .1474  22.50 

Tops    1.5098  22.02  .021 

1.0836  1.2967  18.90  20.46       .018 

Koots     .9436  .4718  18.85  .083 

Tops       .7544  21.18  .105 

.8022  .7783  20.44  20.81        .106 

Roots     .5265  .2632  18.10  .128 


6 

.0000755 

Tops 

.7598 

21.60 

.156 

.6632 

.7125 

20.83 

21.41 

.147 

Roots 

.5889 

.2944 

19.71 

.101 

7 

.0000945 

Tops 

.3339 

23.21 

.374 

.2439 

.2889 

23.60 

23.45 

.294 

Roots 

.1567 

.0783 

18.00 

.982 

8 

.000189 

Tops 

.3267 

22.61 

.673 

.3786 

.3526 

24.80 

23.70 

.480 

Roots 

.1630 

.0815 

21.40 

1 .430 

9 

.000378 

Tops 

.2695 

24.22 

.530 

.2372 

.2533 

22.00 

23.11 

.407 

Roots 

.0696 

.0348 

23.80 

1.53 

3 
.110 

.061 

.205 

.020 

.105 

.151 

.334 


a  to 


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.058 
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1.31 

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.025      

.003 


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.004 


.001 


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Grown  April  14-May  27,  191 


1918] 


Waynick :   Antagonism  and  Cell  Permeability 


205 


l  .  4 


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Fig.   19 

Copper  Sulphate. 

(See  Table  19) 


206 


University  of  California  Publications  in  Agricultural  Sciences       [Vol.  3 


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Waynich:    Antagonism  and  Cell  Permeability 


20^ 


1  .4 


1  .3 


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Fig.  20 

Copper  Sulphate  +  Zinc  Sulphate 

(See  Table  20) 


208 


University  of  California  Publications  in  Agricultural  Sciences       [Vol.  3 


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L918] 


Waynick:    Antagonism  and  Cell  Permeability 


209 


1.4- 


1  .  3 


Fig.  21 

Copper  Sulphate  +  Ferric  Sulphate 

(See  Table  21) 


210 


University  of  California  Publications  in  Agricultural  Sciences        [Vol.3 


Table  22 
Ferric  Sulphate 


No. 

Solution 
Pe2(S04)3 

Dry  Weight 

a 

a 

3 

3 

3 

«  c 

1 

.0000014 

Tops 

2.2399 

17.09 

.107 

.098 

.246 

1.3062 

1.7730 

17.55 

17.32 

.099 

.103 

.113 

.105 

.281 

.263 

Roots 

1.0626 

21.81 

.122 

.763 

.315 

.0000028 

Tops 

.5313 

20.50 

21.81 

.119 

.122 

.070 

.763 

.264 

.315 

2 

1.9198 

1.5226 

1.7210 

20.21 

21.35 

.115 

.117 

.072 

.071 

.201 

.231 

Roots 

.8731 

25.50 

.013 

.145 

.220 

.4850 

.6769 

26.50 

25.85 

.020 

.016 

.158 

.150 

.286 

.253 

3 

.0000070 

Tops 

1.4150 

24.00 

.095 

.393 

1.5247 

1.4698 

22.62 

23.31 

.080 

.087 

.102 

.102 

.395 

.394 

Roots 

.6984 

31.10 

.032 

.263 

.223 

.7624 

.7304 

30.61 

30.85 

.041 

.036 

.224 

.243 

.277 

.250 

4 

.000014 

Tops 

2.3544 

20.57 

.032 

.089 

.358 

2.3461 

2.3502 

18.90 

19.73 

.041 

.036 

.106 

.097 

.378 

.368 

Roots 

1.0361 

27.20 

.022 

.184 

.172 

1.1861 

1.1111 

29.20 

28.31 

.011 

.016 

.250 

.217 

.178 

.175 

5 

.00007 

Tops 

4.2000 

14.80 

.018 

.072 

.236 

3.6033 

3.9016 

15.15 

14.97 

.017 

.017 

.073 

.072 

.268 

.252 

Roots 

.9813 

25.81 

.056 

.134 

.103 

.9168 

.9490 

26.00 

25.90 

.077 

.067 

.118 

.128 

.129 

.116 

Full  Nutrient 

Tops 

1.5682 

18.33 

.293 

.231 

1.5775 

1.5728 

19.40 

18.86 

.312 

.302 

.279 

.255 

Roots 

.9776 

20.40 

.271 

.279 

Grown  April  22-June  3,  1916. 


1918] 


Waynick:    Antagonism  and  Cell  Permeability 


211 


1  .  4 


1  .3 


1  .2 


1  .1 


1.0  - 


0  .9 


0  . 


0.7- 


0  .6 


0  .5 


0  .4 


0  .3 


0  .2 


0  .  1 


\ 


Fig.  22 
Ferric  Sulphate 
(See  Table  22) 


212 


University  of  California  Publications  in  Agricultural  Sciences       [Vol.3 


Table  23 

Zinc   Sulphate 

No. 

Solution 
ZnS04 

Dry  Weight 

3 

M 
1  « 

K       ° 

3 

bfl 

a 

h 

1 

.00000767 

Tops 

.6718 

23.00 

.324 

.7541 

.7129 

24.04 

23.52 

.341 

Eoots 

.6138 

.3069 

17.88 

.191 

2 

.0000131 

Tops 

.7164 

25.30 

.326 

.8350 

.7757 

24.65 

25.07 

.284 

Roots 

.6009 

.3004 

17.20 

.187 

3 

.0000395 

Tons 

.6142 

21.19 

.372 

.8607 

.7374 

22.50 

21.84 

.316 

Roots 

.5864 

.2932 

15.90 

.166 

4 

.000153 

Tops 

.5109 

21.90 

.372 

.3800 

.4454 

19.40 

20.65 

.437 

Roots 

.4628 

.2314 

20.50 

.128 

5 

.000395 

Tops 

.8208 
Lost 

.4104 

20.38 

20.38 

.200 

Roots 

.4432 

.2216 

22.30 

.053 

Ful 

1  Nutrient 

Tops 

1.0992 

20.17 

.310 

1.0750 

1.0872 

19.12 

18.69 

.297 

Roots 

.8120 

20.00 

.271 

.332 


.305 


.344 


.404 


.200 


.303 


C   60 


.374 
.560 


.810 

.268 
.228 
.233 


.46^ 


.0622 

.0587 


.310 

.121 

.386 

.348 

.101 

.358 

.101 

,324 

.0708 

.320 

.322 

.0644 

.886 

.660 

.097 

.760 

.710 

.981 

.089 

.423 

.074 

321 

.073 

.321 

.073 

.01' 


.224 


Grown  January  24-March  6,  1916. 


1918] 


Waynick:   Antagonism  and  Cell  Permeability 


213 


1  .0 


0  .9 


0  .  8 


0.7- 


0.6- 


0  .5 


0.4- 


/\ 


0  .3 


0  .2 


0.1- 


\ 


\ 


Fig.  23 
Zinc  Sulphate 
(See  Table  23) 


214 


University  of  California  Publications  in  Agricultural  Sciences       [Vol.  3 


Table  24 
Zinc  Sulphate  +  Ferric  Sulphate 


Solution 


Xo.      ZnS04       Fe2(SOJ3 
1      .0000019  .0000035 


Dry  Weight 

Tops     1.0594 

1.5294 

Eoots  .10444 


2  .0000038  .000005      Tops    1.5364 

1.4864 
Boots  1.0716 

3  .0000057  .000007      Tops    1.2300 

1.6150 
Eoots  1.1250 


1.2944 
.5222 


18.50 

18.70     18.60 

29.80 


18.60 
1.5114     16.10     1' 
.5258     31.00 


6      .0000379  .000070 


7     .000076     .00014       Tops    1.6164 

1.7228 
Eoots  1.7611 


8      .000152     .00028 


Tops       .9850  18.35 

1.0268     1.0029     20.00 

Eoots     .6987       .3493     27.30 


.35 


18.10 
1.4225  16.90  17.00 
.5625  32.30 


4  .0000076  .000014  Tops  2.3088  16.40 

2.5100  2.4094  17.00  16.70 

Eoots  1.5623  .7812  28.00 

5  .0000152  .000028  Tops  2.3429  15.70 

2.0129  2.1774  15.70  15.70 

Eoots  1.5133  .7566  30.30 


Tops  2.0533        18.10 

1.5544  1.8038  16.20  17.15 
Eoots  1.4194   .7097  25.30 


18.40 
1.6696  19.70  19.05 
.8801  27.00 


19.17 


On  ~ 

.096 
.098 
.045 

.149 
.170 
.042 

.172 
.124 
.068 

.083 

.098 
.040 

.154 
.147 
.050 

.160 
.160 
.025 

.157 
.144 
.019 

.179 
.176 
.006 


.097 


159 


.140 


.090 


.150 


.160 


.150 


.177 


O) 

Ph 

3 

.215 
,230 
.149 

.222 

.031 
.029 
.037 

.030 

.262 
.231 

.184 

.247 

.036 
.030 
.082 

.033 

.288 
.277 
.130 

.282 

.076 
.064 
.010 

.070 

.110 
.111 
,139 

.110 

.065 
.062 
.061 

.063 

.151 
.158 
.103 

.154 

.052 
.046 
.069 

.049 

.122 
.151 
.243 

.136 

.042 
.049 
.141 

.045 

.201 
.317 

.0412 
.0412 
.128 

.043 

.219 
.282 
,367 

.250 

.045 
.042 

.043 

Grown  April  24-June  6,  1916. 


1918] 


Waynick:   Antagonism  and  Cell  Permeability 


215 


0  .5 


0  .4 


0.3- 


-  -  -  -^K 


XV 


/       v 

/  \ 

/  \ 


\ 


o  .1 


^:\>< 


Fig.  24 

Zinc  Sulphate  +  Ferric  Sulphate 

(See  Table  24) 


216 


University  of  California  Publications  in  Agricultural  Sciences       [Vol.  3 


Table  25 
Mercuric  Chloride  4-  Ferric  Sulphate 


Solution 

A 


SO 

Pm 


No.         HgCl2       Fe2(S04)3    Dry  Weight  ^  Pn  °  g 

1      .0000047  .0000035    Tops    .2039  22.40  .233 

.2868  .2553  16.70  19.55     .179     .206 

Roots  .0518  .0259  18.70  .840 


2  .0000094  .0000070  Tops 

Roots 

3  .0000189  .00005        Tops 

Roots 

4  .000047     .00014       Tops 

Roots 

5  .000094     .0007  Tops 

Roots 

6  .000189     .00105        Tops 

Roots 

7  .000378     .00210        Tops 

Roots 


.2569 
.3579 
.0869 

.2042 
.2996 
.0896 

.2362 
.2300 
.0596 

.2396 
Lost 
.0237 

.2288 
.2988 
.0784 

.2184 
.4272 
.0496 


21.75 
.3074     21.90     21.82 
.0434     18.90 

16.70 
.2519     19.40     18.05 
.0448     18.13 


17.36 


17.32 
.2331  17.41 
.0298     19.01 

16.10 

.2396      

.0118     15.50 

16.70 
.2638     17.20     16.95 
.0392     21.30 


.123 

.138     .230 
.222 

.172 

.231     .201 

.271 

.192 

.183     .187 

.221 

.420 
16.10      420 


.151 

.201 

.131     .166 


15.61 
.3228  17.40 
.0248     18.10 


16.50 


.190 
.102 
.232 


.146 


££  g 

Ph  S 

.723 

.513     .618 
2.670 


.327 
.515 
2.95 


.421 


.371 

.412     .391 

.312 


.416 

.511 

2.110 

.100 


.100 


4.580 


.731 

.807     .769 
2.170 


.895 
.550 

.738 


.463        ft 


.722 


o 
a 

o 


.101 
.157 

.222 


.129 


Grown  March  22-May  3,  1916. 


1918] 


Waynick:    Antagonism  and  Cell  Permeability 


21' 


1  .  0 


0  .9 


0.8  - 


0.6- 


0  .  5 


0  .  4 


0  .  3 


0  .  2 


0.1- 


Fig.  25 

Mercuric  Chloride  +  Ferric  Sulphate 

(See  Table  25) 


218  University  of  California  Publications  in  Agricultural  Sciences       [Vol.3 


Table  26 
Mercuric  Chloride 

O  ( 

hi)  i 


Solution  .as  S*S  a  £*£ 


c3 

Xo.         HgCl2             Dry  Weight  ^              Ph  ~  S              P^ 

1  .0000135      Tops     .4904  17.20  .141 

.4967  .4935       16.71  16.95       .098       .120 

Eoots  .1493  .0746       19.31  .478 

2  .000066        Tops    .2200  15.70  .332 

.2236  .2218       14.24  14.97       .380       .356 

Eoots  .0239  .0119       10.50  4.31 

3  .000135        Tops    .1514  9.12  .421 

.1421  .1467          8.95  9.03       .399       .410 

Roots    


c  fan 

a 

3 

.91 

.248 

.91 

.248 

.893 

.296 

1.69 

.010 

1.77 

1.73 

.017 

.013 

.775 

.013 

1.23 

.012 

1.33 

1.28 

.013 

.012 

Grown  March   14-April  24,  1916. 


1918] 


WaynicJc:    Antagonism  and  Cell  Permeability 


219 


1  .4- 


1  .3 


1  .  2 


1  .1 


1  .  0 


0  .9 


0  .  6 


0.5- 


0.4- 


0  .3 


0  .2 


0  .1 


/ 


./ 


\ 


I  Is 


Fig.  26 

Mercuric  Chloride 

(See  Table  26) 


EXPLANATION    OF    PLATES 

PLATE  13 

Appearance  of  plants  as  mounted  in  corks  at 
expiration  of  the  six  weeks'  growing  period. 


[220] 


UNIV.    CALIF.    PUBL.    AGR.    SCI.    VOL.    3 


[WAYNICK]    PLATE    13 


1 


PLATE  14 

No.     1.  .24  M.  MgCl2  .004  M.  CaCl2 

No.     2.  .24  M.  MgCl2  .01  M.  CaCL 

No.    3.  .24  M.  MgCl2  .02  M.  CaCl2 

No.     4.  .24  M.  MgCl2  .04  M.  CaCl2 

No.     5.  .24  M.  MgCl2  .06  M.  CaCl2 

No.     6.  .24  M.  MgCL  .08  M.  CaCl2 

No.     7.  .24  M.  MgCL  .10  M.  CaCl2 

No.     8.  .24  M.  MgCl2  .12  M.  CaCL 

No.     9.  .24  M.  MgCl2\l6  M.  CaCL 

No.  10.  .24  M.  MgCL  .20  M.  CaCL 

No.  11.  .24  M.  MgCl2  .24  M.  CaCl2 

No.  12.  .24  M.  MgCL  .30  M.  CaCL 

No.  13.  .24  M. 

Control. 


|  222 


UNIV.    CALIF.    PUBL.    AGR.    SCI.    VOL.    3 


[WAYNICK]    PLATE     14 


; 


\    w 


- .    J 

■ill 

1 

PLATE  15 

No.    1.  .30  M.  MgCl2  .004  M.  CaCL 

No.    2.  .30  M.  MgCl2  .01  M.  CaCl2 

No.    3.  .30  M.  MgCl2  .02  M.  CaCl2 

No.    4.  .30  M.  MgCl2  .04  M.  CaCl2 

No.    5.  .30  M.  MgCl2  .06  M.  CaCl2 

No.    6.  .30  M.  MgCl2  .08  M.  CaCl2 

No.    7.  .30  M.  MgCl2  .10  M.  CaCl2 

No.    8.  .30  M.  MgCl2  .12  M.  CaCl2 

No.    9.  .30  M.  MgCl2  .16  M.  CaCl2 

No.  10.  .30  M.  MgCl2  .20  M.  CaCl2 

No.  11.  .30  M.  MgCl2  .24  M.  CaCl2 

No.  12.  .30  M.  MgCl2  .30  M.  CaCl2 
Control. 


224] 


UNIV.    CALIF.    PUBL.    AGR.    SCI.    VOL.    3 


[WAYNICKJ     PLATE     15 


V    4 


PLATE  1( 

No. 

1. 

.04  M. 

KC1 

No. 

2. 

.06  M. 

KC1 

No. 

3. 

.08  M. 

KC1 

No. 

4. 

.10  M. 

KC1 

No. 

5. 

.12  M. 

KC1 

No. 

6. 

.14  M. 

KC1 

No. 

7. 

.16  M. 

KC1 

[  220  | 


s 


;<    -     i. 


fW 


PLATE  17 

No. 

1. 

.00331   M. 

A1C1, 

No. 

2. 

.000331  M. 

A1C13 

No. 

3. 

.000132  M. 

A1C1, 

No. 

4. 

.000066  M. 

Aid, 

No. 

5. 

.000033  M. 

AlClj 

No. 

6. 

.0000165  M. 

A1C13 

No. 

7. 

.0000033  M. 
Control. 

AICI3 

228  I 


-J 


PLATE  18 

No.  1.  .0168      M.  FeCl3  .20  M.  MgCl2 

No.  2.  .0058      M.  FeCl3  .20  M.  MgCl2 

No.  3.  .00356    M.  FeCl3  .20  M.  MgCL 

No.  4.  .00142    M.  FeCl3  .20  M.  MgCL 

No.  5.  .000712  M.  FeCl3  .20  M.  MgCl2 

No.  6.  .000352  M.  FeCl3  .20  M.  MgCL 

No.  7.  .000168  M.  FeCl3  .20  M.  MgCL 

No.  8.  .000089  M.  FeCl3  .20  M.  MgCL 
Control. 


I'M) 


PLATE  19 

No.  1.  .00331      M.  A1C13  .20  M.  CaCL 

No.  2.  .000331    M.  A1C13  .20  M.  CaCL 

No.  3.  .000132    M.  A1C13  .20  M.  CaCL 

No.  4.  .0000662  M.  A1C13  .20  M.  CaCL 

No.  5.  .0000331  M.  A1C13  .20  M.  CaCl2 

No.  6.  .0000165  M.  A1C13  .20  M.  CaCl2 

No.  7.  .0000033  M.  A1C13  .20  M.  CaCl2 
Control. 


[232] 


PLATE  20 

No.  1.     .000094    M.  CuCl2  .00082  M.  FeCl3 

No.  2.     .000067    M.  CuCl2  .000058  M.  FeCl3 

No.  3.     .000047    M.  CuCl2  .000042  M.  FeCl3 

No.  4.     .000028    M.  CuCL  .000026  M.  FeCl3 
No.  5.     .0000094  M.  CuCL  .0000089  M.  FeCl3 
Control. 


[234] 


^ 


PLATE  21 

No. 

1. 

.000378     M. 

CuS04 

No. 

2. 

.000189     M. 

CuS04 

No. 

3. 

.0000945  M. 

CuS04 

No. 

4. 

.0000755  M. 

CuS04 

No. 

5. 

.0000567  M. 

CuS04 

No. 

6. 

.0000378  M. 

CuS04 

No. 

7. 

.0000188  M. 

CuS04 

No. 

8. 

.0000094  M. 

CuS04 

No. 

9. 

.0000048  M. 
Control. 

CuS04 

[236] 


PLATE  22 

No.  1.  .0000047  M.  CuSo4  .0000035  M.  Fe2  (S04)3 

No.  2.  .0000094  M.  CuSo4  .0000070  M.  Fe2  (S04)8 

No.  3.  .0000142  M.  CuSo4  .0000105  M.  F'e2  (S04)3 

No.  4.  .0000189  M.  CuSo4  .000014     M.  Fe2  (S04)3 

No.  5.  .0000378  M.  CuSo4  .000028     M.  Fe2  (S04)3 

No.  6.  .000094    M.  CuSo4  .000070     M.  Fe2  (S04)3 

No.  7.  .000142     M.  CuSo4  .000105     M.  Fe2  (S04)3 

No.  8.  .000189     M.  CuSo4  .00014       M.  Fe2  (S04)3 

No.  9.  .00058       M.  CuSo4  .00028       M.  Fe2  (S04)3 
Control. 


[238] 


PLATE  23 

No. 

1. 

.0000014  M.  Fe2 

(S04)3 

No. 

2. 

.0000028  M.  Fe2 

(S04)3 

No. 

3. 

.0000070  M.  Fe2 

(S04)3 

No. 

4. 

.000014     M.  Fe2 

(S04)3 

No. 

5. 

.00007       M.  Fe2 
Control. 

(S04)3 

[240] 


PLATE  24 

STo.  1. 
STo.  2. 
STo.  3. 

.000135     M. 
.000066     M. 
.0000135  M. 
Control. 

HgCl2 
HgCl2 
HgCL 

[242] 


UNIV.    CALIF.    PUBL.    AGR.    SCI.    VOL.    3 


[WAYNICK]    PLATE    24 


i  / 


/// 


